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Full text of "Bulletin of the British Museum (Natural History) Zoo Supp"

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BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



ZOOLOGY 
VOL. V 
195 7-1960 




PRINTED BY ORDER OF THE TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

LONDON: i960 



DATES OF PUBLICATION OF THE PARTS 



No. i. 
No. 2. 
No. 3. 
No. 4. 
No. 5. 
No. 6. 
No. 7. 
No. 8. 
No. 9. 
No. 10. 



26 February 1957 

21 March 1957 

30 July 1957 

12 October 1957 

26 August 1958 

24 January 1959 

24 February 1959 

24 February 1959 

24 February 1959 

22 January i960 



PRINTED IN 

GREAT BRITAIN 

AT THE 

BARTHOLOMEW PRESS 

DORKING 

BY 

ADLARD AND SON, LTD. 



CONTENTS 



ZOOLOGY VOLUME 5 PAGE 

No. I. Studies on the structure and taxonomy of Bulinus jousseaumei 

(Dautzenberg). By C. A. Wright (Pis. 1-2) 1 

No. 2. On Spelaeogriphus, a new cavernicolous crustacean from South Africa. 

By Isabella Gordon 29 

No. 3. The pelecaniform characters of the skeleton of the Shoe-bill Stork, 

Balaeniceps rex. By Patricia A. Cottam (PI. 3) 49 

No. 4. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) 
Part II : H. sauvagei (Pfeffer), H. prodromus Trewavas, H. granti 
Blgr., and H. xenognathus, sp. n. By P. H. Greenwood (PI. 4) 76 

No. 5. A revision of the genera Nidalia and Bellonella, with an emendation of 
nomenclature and taxonomic definitions for the family Nidaliidae 
(Octocorallia, Alcyonacea). By Huzio Utinomi 99 

No. 6. Ear plug laminations in relation to the age composition of a popula- 
tion of fin whales (Balaenoptera physalus). By P. E. Purves and 

M. D. MOUNTFORD (Pis. 5-6) 123 

No. 7. The monotypic genera of cichlid fishes in Lake Victoria Part II and a 
revision of the Lake Victoria Haplochromis species (Pisces Cichlidae) 
Part III. By P. H. Greenwood 163 

No. 8. The Rosaura Expedition 1937-38 Chaetognatha. By John S. Colman 219 

No. 9. A North Bornean pygmy squirrel, Glyphotes simus Thomas, and its 

relationships. By J. E. Hill (Pis. 7-8) 255 

No. Io. Revision of the world species of Aplysia (Gastropoda, Opistho- 

branchia). By N. B. Eales 267 

Index to Volume 5 405 




STUDIES ON THE 

STRUCTURE AND TAXONOMY 

OF BULINUS JOUSSEAUMEI 

(DAUTZENBERG) 



C. A. WRIGHT 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. i 

LONDON: 1957 



STUDIES ON THE STRUCTURE AND 

TAXONOMY OF BULINUS JOUSSEAUMEI 

(DAUTZENBERG) 



BY 

C. A. WRIGHT 



Pp. 1-28 ; Plates 1-2 ; 41 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 1 

LONDON: 1957 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5, No. 1 of the Zoological series. 




PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued February 1957 Price Ten Shillings 



STUDIES ON THE STRUCTURE AND 
TAXONOMY OF BULINUS JOUSSEAUMEI 
(DAUTZENBERG) 

By C. A. WRIGHT 

SYNOPSIS 
Many recent workers on the freshwater mollusca of Africa have tended to present synonymies 
based more on the literature and on personal opinions than on a study of the animals themselves. 
This is particularly true of those gastropods of medical importance which act as intermediate 
hosts for flukes parasitizing human beings. This paper is an attempt to establish the relation- 
ships of one such snail. 

HISTORICAL 
Dautzenberg (1890) described Bulinus jousseaumei as a member of the genus 
Isidora and stated that it belonged to the /. contorta group with a strong affinity to 
/. natalensis. Pilsbry & Bequaert (1927) list B. jousseaumei in the sub-genus 
Bulinus s. str. and Amberson and Schwarz (1953) place it in the synonomy of B. 
truncatus. I have shown that both on conchological and anatomical grounds the 
species is properly placed in the sub-genus Physopsis (Wright 1956). Smithers 
(1956) has shown that in parts of the Gambia Protectorate this snail is an important 
vector of the human blood-fluke, Schistosoma haematobium. From personal 
observations I am sure that it is also the vector implicated in the Casamance 
Province of Senegal by Deschiens (1951) under the name Bulinus trigonus, a 
species characteristic of some lakes in East Africa. 

MATERIAL AND METHODS 

The material of B. jousseaumei from the Gambia and Senegal used in this work was 
collected personally, in collaboration with S. R. Smithers. Material from other 
localities has mostly been seen in the collection of the British Museum (Natural 
History) or has been sent to me by other workers. 

Shell measurements were made with an eyepiece micrometer in a binocular 
dissecting microscope, the shells were held steady in a horizontal position by means 
of a piece of plasticine on the microscope stage. Radula preparations were made 
following the rapid methods recommended by Meeuse (1950). 

THE SHELL 

The original description of the shell by Dautzenberg is very adequate and my re- 
description (Wright, 1956) adds little to the original, apart from noting a further 
colour variation from light yellowish-straw to dark reddish-brown, and amplifying 

ZOOL. 5, I. 



4 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUME1 

the details of the ornamentation. Since reference will be made later to the sculpture 
my previous observations are repeated here. There is a regular punctate pattern 
on the nuclear whorl and the rest of the shell is covered with fine growth lines 
superimposed on which is a delicate pattern of short, wavy, vertical lines. It is 
surprising that Dautzenberg, who must have been acquainted with Physopsis, was 
so definite about the affinities of B. jousseaumei in spite of his mention of the twisted 
base of the columella in his description. 

One of the greatest difficulties in the description of gastropods is the definition of 
size. The old workers when describing a new species usually gave the dimensions 
of the specimen (often the largest) which they had selected as the holotype. Rarely, 
they gave the dimensions of the largest and the smallest specimens available to 
them. The dangers of this approach to the problem in an animal which grows 
steadily without a well-defined adult phase are obvious. Size ranges are valueless 
since they give no idea of distribution, and ranges given with a mean and standard 
deviation are not of much use unless the growth stages represented in the sample 
are known. The ratios of certain measurements of the shell are of more value but 
these may vary with shell length. For instance Peters (1938) has shown that the 
ratio shell length /aperture length increases with shell length in Lymnaea palustris. 
Hubendick (1951) using smaller samples suggests that there is no significant change 
in the ratio aperture length/shell length x 100 in Lymnaea peregra. 

In order to investigate the relationships of B. jousseaumei it has been found 
necessary to analyse the size composition of various samples. The largest of these 
samples was collected at a washing place in the Simoto Bolon near the village of 
Diabugu Basilla, Upper River Division, Gambia on the 4th March, 1955. Two- 
hundred and thirty-eight shells were measured and spirit material dissected to try to 
determine an approximate correlation between shell size and the attainment of the 
" adult " condition. The results obtained from a study of this sample appear to be 
characteristic for the other samples collected in the Gambia at the same time. The 
time of year at which a sample is collected is of great importance since it is almost 
certain that the population early in the wet season contains a much higher proportion 
of juveniles than it would some months after the end of the rains, the time when the 
material under consideration was collected. Text-fig. 1 is a size frequency histogram 
of this sample, based on shell length. It is obvious that the greatest frequency 
occurs in the 8-0-9-0 mm. shell length group and the mode of the graph is at the 8-5 
mm. level. The graph is, however, asymmetrical due to an overweighting of the 
lower size groups, the mean shell length of the sample is therefore less than the modal 
length. The graph does not provide any indication of whether the mode is due to 
large numbers of adults or juveniles and this can only be determined by anatomical 
study. It will be shown later that it is in the size group 6-0-7-0 mm. shell length 
that the accessory reproductive glands reach a fully functional stage of their develop- 
ment and this is taken as the beginning of the adult phase of life. If the curve show- 
ing distribution on the upper side of the mode in Text-fig. 1 is drawn and its mirror 
image is reproduced on the low side a normal distribution curve is formed, the lower 
end of which coincides well with the known onset of the adult phase. If the 
frequency distribution is plotted on arithmetical probability paper following the 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 5 



1 ' 



2 3 4 5 * » • 

SHELL LENGTH IN MILLIMETRES 



1.4 






.. 




2.5 
2.4 








• 


2.) 






" • . " "*.'• 




L/MB 






'■• '* .;>:•: .:':' .• 




2.1 






• 




1 






• 




l.f 








2 


1-4 










I.J 
L/ML 








. 


1.2 


; 


: • • 


,.«;-^lffev'' 


• 


bl 




.v. 




3 



1 4 5 • 7 « 

SHELL LENGTH IN MILLIMETRES 



Fig. 1. Size-frequency histogram for a population of B. jousseaumei from 

the Gambia. 
Fig. 2. Graph of ratio shell length /aperture width against shell length for 

Gambian population of B. jousseaumei. 

Fig. 3. Graph of ratio shell length /aperture length against shell length 

for the same population. 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 



method of Harding (1949) a curve is obtained which appears to be too complex to 
be analysed by the usual methods. This may in part be due to faulty sampling but 
is also undoubtedly caused by the complicated structure of a population of fresh 
water snails. There may be present the representatives of several successive 
generations, a few senescent survivors of previous generations, the adults of the 
present and the progeny of these adults. If the breeding season is continuous then 
a smooth curve might be expected but there is reason to believe that breeding is 
somewhat spasmodic in the bulinids and this might help to account for the complex 
composition of the population. Since the anatomical findings confirm that the mode 
of the histogram in Text-fig. 1 is in fact the mean of the adult part of the population 
it seems justifiable to accept 8-5 mm. as the mean shell length of the adults in this 
sample. An indication of the proportion of juveniles in the whole sample is given 
by the index of skewness, (mean-mode) /standard deviation. Having obtained a 
rough mean length for the adults in the population the adult means for the other 
dimensions may be calculated by increasing the population mean by the same 
percentage as that by which the mean population length differs from the adult mean 
length. The following table gives the mean shell dimensions of the population in 
millimetres and their standard deviations together with the approximate adult 
dimensions and the maximum sizes observed. 



Aperture 



Whole population 
Adults . 
Maximum 



Length 

A 

( \ 

Mean S. D. 

7-84 ±1-52 

8-5 

n-8 



Max. diam. 



Mean S. D. 

6 • 00 4: 1 • 1 2 
6-50 — 

8-o 




Aperture 
width 



Mean S. D. 



3-37 
3'6 5 



±o-66 



4'5 



Passing from the absolute dimensions of the shells to the ratios of some of these 
dimensions to one another, one is faced with a number of these proportions from 
which to choose the most useful. Hubendick (195 1) pointed out that the selection 
of measurements in any particular case should be made with advance knowledge of 
the variation to be studied. In this study of B. jousseaumei the object of interest is 
not so much phenotypic or genotypic variation as changes in form during growth. 
With this in mind the two ratios chosen are those of shell length /aperture length 
(1/ml) and shell length /aperture width (1/mb). The first of these in this type of shell 
gives an indication of the exsertion of the spire for if the spire is strongly depressed 
the ratio will approach unity and will increase with the exsertion of the upper whorls, 
or, more properly, the descent of the body whorl. The second of the two ratios 
expresses the relationship between the increase in diameter of the body whorl with 
increasing shell length. A similar result might easily be obtained by using the ratio 
shell length /shell diameter but with the technique of measurement employed the 
width of the aperture was more accurately obtainable than the maximum diameter 
of the shell. Text-figs. 2 and 3 show these two ratios plotted against shell length ; 
in both it can be seen that there is an increase in the value of the ratio with increasing 
shell length. The change in the values is so slight that the means of the ratios 
provide a good index of shell form, The graph 1/ml plotted against shell length 



THE STRUCTURE AND TAXONOMY OF BlfLINUS JOUSSEAUMEI 7 

(Text-fig. 3) shows an interesting feature. For small shell lengths the graph is 
practically a straight line parallel to the horizontal axis and the upward slope only 
becomes apparent after the 6*5 mm. shell length mark. As pointed out earlier, this 
is the approximate shell length at which adult anatomical characters become 
developed and the change in form of the shell at this point is, presumably, a reflexion 
of the anatomical changes. The actual change in form which the shell exhibits may 
be explained thus : in its early stages the spire is completely depressed, subsequent 
whorls being added around the preceding ones ; at the onset of maturity the accessory 
genital glands increase in size very rapidly necessitating an increase in shell volume 
to accommodate them and this is achieved by the body-whorl moving downward in 
relation to its predecessor, giving an increase in shell length without a corresponding 
increase in aperture length. 

The means and standard deviations of these two ratios in the population under 
consideration are 

1/ml mean 1-17 s.d. ^ 0-08 

1/mb mean 2.31 s.d. ± 0'i43 

THE MANTLE 

The mantle markings of B. jousseaumei from the Gambia consist of patterns of 
small black spots and patches scattered irregularly over a light grey field. The 
spots often appear dark grey rather than black owing to the pigment granules of 
which they are composed being only loose aggregations rather than dense 
concentrations. 

The nephridial ridge on the underside of the mantle may be well developed or 
almost absent. It is often present only on the distal end of the kidney. The inter- 
mediate ridge is almost always well developed and is about equal in length to the 
kidney. An examination of transverse sections of the mantle has failed to show 
the presence of ciliated epithelium on this intermediate ridge (described for B. 
africana by Hubendick, 1948) but this may be due to the method of fixation employed. 

THE RADULA 

The radula is in no way remarkable. The number of tricuspid laterals varies 
from six to eight in each transverse half-row. This number does not appear to 
change with the age of the snail, but the number of marginal teeth does appear to 
increase in older specimens. It is perhaps worthwhile recording here that the first 
lateral tooth is tricuspid (Text-fig. 4). Dupuis and Putzeys (1923) mention as the 
only real difference between Physopsis and Isidora that the first lateral in the former 
group is bicuspid while in the latter it is tricuspid. The endocone of the first lateral 
is often difficult to observe and it is doubtless this fact that gave rise to the erroneous 
statement of these authors. 

CENTRAL NERVOUS SYSTEM 

The central nervous system in the Planorbidae shows little variation between the 
various genera. The connectives between the ganglia are relatively long in B. 
jousseaumei, but since the material on which these anatomical observations are 



8 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

based was all narcotized and well extended such details may not be strictly compar- 
able with those of other workers (Text-fig. 5). 

The penial nerve appears to be a composite structure with its main source in the 
left cerebral ganglion ; a number of other fibres running with the main component 
arise from the left pedal ganglion. The cerebral part of the nerve branches off from 
a larger trunk arising near the origin of the left cerebro-buccal connective and passing 
forwards giving off finer branches to the sides of the head and lips. It is probable 
that the nerves arising from the cerebral ganglia have a primarily sensory function, 
while those from the pedal ganglia are mainly motor. This would mean that the 
composite nature of the penial nerve provides both sensory and motor innervation 
for the copulatory organ. The principal nerve arising from the dorsal surface of 
each of the cerebral ganglia runs to the tentacle, its associated lobe and the eye on 
the same side of the body as that from which it arises. The pedal ganglia send 
several large nerves down into the foot and running back from the two buccal 
ganglia is a pair of fine nerves, one on either side of the oesophagus. The three 
visceral ganglia send nerves to the organs in the visceral mass and one large trunk 
arising from the left visceral ganglion passes to the anal lobe and pseudo-branch, 
while a similar large trunk from the right visceral ganglion passes upwards to the 
mantle. Contributory evidence as to the sensory nature of the cerebral ganglia is 
obtained from the otocyst which, although partially embedded in the posterior side 
of the pedal ganglion appears to be innervated solely from the cerebral ganglion 
above (Text-fig. 6). 

BLOOD CIRCULATORY SYSTEM 

The heart lies within its extremely delicate pericardium on the mantle close to the 
proximal end of the kidney, above the point at which the oesophagus passes into 
the crop. The auricle receives blood from the very large vein running along the 
anterior edge of the kidney. The aorta leaving the ventricle is variable in length, 
it may pass right over the intestinal loop which curves round the gizzard before 
dividing (Text-fig. 7) or it may divide so far back that the ventricle has a bifid 
appearance. In either case, the lesser of the two branches follows the intestine on 
its course round the gizzard while the major branch passes upward over the posterior 
edge of the gizzard. As it passes over the space between the intestine and gizzard a 
very large artery passes down between these two organs and divides almost at once, 
one branch going to the crop and distal side of the gizzard, the other to the accessory 
genital glands and the head cavity. A short distance after this division of the main 
branch from the aorta a smaller branch is given off to supply the stomach and 
proximal part of the gizzard and the rest of the vessel continues up the side of the 
gizzard, past the point at which the digestive gland opens from the intestine, and 
then, giving off a number of side branches into the digestive gland, follows the path 
of the intestine as it loops up into the upper whorls of the body. The principal 
artery to the head follows beneath the oesophagus to the circum-oesophageal nerve 
ring where vessels supplying the ganglia are given off ; then, after passing through 
the nerve ring it divides. One branch passes vertically downwards as the pedal 
artery and the other continues forward to the underside of the buccal mass where the 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 9 






Fig. 4. Radula teeth of B. jousseaumei from Gambia. 

Fig. 5. Central nervous system of B. jousseaumei. 

Fig. 6. Right lateral view of central nervous system of B. jousseaumei to show 

the otocyst and its innervation 
Fig. 7. Heart and principal arteries of B. jousseaumei. 
(Figs. 5, 6 8c 7 to same scale.) 
zool. 5, 1 I; 



io THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

vessel dilates before breaking up into fine branches supplying the muscles of the mass 
and the sides of the head and lips. A fine lateral branch which leaves the main 
trunk in the region of the nerve ring serves the penial complex. It runs parallel with 
the penial nerve and appears to enter the complex at the junction of the penis sheath 
and preputium. 

ALIMENTARY SYSTEM 

The digestive tract in the Planorbidae is so well known and subject to so little 
variation that it is not necessary to enter into a full description here. 

REPRODUCTIVE SYSTEM 

The gross morphology of the genital tract of B. jousseanmei has already been 
described (Wright, 1956). It is intended here to consider the histology of the tract 
and its development. Hubendick (1948 a & b) has described the anatomy and 
histology of the male copulatory organs of several species of Bulinus. Larambergue 
(1939) has described fully the reproductive anatomy and histology of Bulinus 
truncatus. Abdel-Malek (1954 a & b) has given detailed accounts of the morphology 
and histology of the genital organs of two Planorbids Helisoma trivolvis (Say) and 
Biomphalaria boissyi and comparisons between all of these and Bulinus jousseaumei 
will be made. 

Histologically no real differences were found between Abdel-Malek's description of 
the ovotestis in Biomphalaria and Helisoma and that in the present species. The 
acini are enveloped in " Ancel's layer " of thin connective tissue and the germinal 
epithelium within this layer appears to line only the lower parts of the acini. 
Heavily pigmented connective tissue is largely confined to the layer covering the 
top of the organ. In the adult snail all stages of spermatogenesis and oogenesis can 
be observed in the same acinus. Young oocytes and spermatids are largely confined 
to the lower parts of the acini and the upper parts are occupied by maturing ova, as 
many as six having been seen in a single acinus as compared with two to three reported 
by Abdel-Malek in Helisoma trivolvis. The mature ova are enclosed within a follicular 
membrane made up of nurse cells and connective tissue. Mature spermatozoa 
are more or less ubiquitous in the acini, either free in the lumen or attached by their 
anterior ends to basal " Sertoli " cells. The motility of these basal cells is shown 
in that they may be found on the outer wall of the follicles of maturing ova, a position 
that could only be reached by their independent locomotion. The hermaphrodite 
duct is lined with an epithelium of cuboidal cells bearing short cilia and the duct is 
sheathed in a thin layer of connective tissue (Text-fig. 8). The epithelium lining 
the seminal vesicles is similar to that in the hermaphrodite duct but the median 

Transverse section of hermaphrodite duct of B. jousseaumei. 

Transverse section of first part of sperm duct of B. jousseaumei 

Transverse section of sperm duct of B. jousseaumei. 

Epithelial lining of prostate tubule of B. jousseaumei. 

Transverse section of vas deferens within body wall of B. jousseaumei. 

Transverse section of vas deferens in head cavity of B. jousseaumei. 



Fig. 


8. 


Fig. 


9- 


Fig. 


10. 


Fig. 


11. 


Fig. 


12. 


Fig. 


13- 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI n 




12 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

nuclei are a little larger and the cytoplasm is more granular. The separation of the 
male and female ducts from the common duct occurs at a point well embedded in the 
base of the albumen gland. The sperm duct at its source is narrow with a small 
lumen lined by short columnar epithelial cells, ciliated, and with median to basal 
nuclei and finely granular cytoplasm (Text-fig. 9). This part of the duct is quite 
short and it soon becomes considerably larger in diameter although the size of the 
lumen does not increase greatly. The epithelium lining the duct loses its cilia and 
the cells are of a much taller columnar type with median nuclei, larger than those in 
the earlier part of the duct, and the cytoplasm is filled with large, eosinophilic, 
retractile granules. A few wedge-shaped cells occur between the apical ends of the 
columnar type and these do not contain the eosinophilic retractile granules (Text-fig. 
10). The sperm duct retains this histological structure right up to its entry into the 
prostate gland. Hubendick (1948a) in discussing the structure of the prostate in 
Bulinus has shown that in B. inflatus, an Australian species, the tubules of the 
prostate open individually into the vas deferens which enters the gland at the 
proximal end and leaves it distally, a structure similar to that in Physa. In B. 
jousseaumei however the prostate is a discoidal structure, flattened on one side and 
convex on the other. The sperm duct enters the gland almost in the centre of the 
flattened surface and the vas deferens leaves it also almost in the centre. There is 
therefore a central point at which the tubules of the prostate discharge their secretion 
into the male duct and it is at this point that the histological structure of the male 
duct changes from the typical form of the sperm duct to that of the vas deferens. 
The tubules of the prostate gland are close-packed, each is ensheathed in a thin-layer 
of connective tissue and is lined by large secretory cells with large basal nuclei with 
granular contents. The cytoplasm of these cells is eosinophilic and not granular 
(Text-fig. 11). There are some basophilic cells near the blind ends of the tubules. 
As the vas deferens leaves the prostate it is lined with sparsely ciliated epithelium 
of rather flattened cuboidal cells with central nuclei containing sparse chromatin 
granules. This epithelium is surrounded by a thin layer of circular muscle. As the 
duct proceeds on its course there is an increase both in the ciliation of the lining 
epithelium and in the thickness of the circular muscle layer. Where the duct passes 
through the body wall the lumen is large, the cells of the epithelial lining are wider 
than high, the cilia are plentiful but not long and the muscle layer is thick (Text-fig. 
12). After leaving the body wall and entering the head cavity the structure of the 
vas deferens changes slightly. The lumen becomes more restricted, the epithelial 
lining more columnar, with basal nuclei, still with few chromatin granules in the 
clear nucleoplasm, the cilia become longer and the muscle layer thicker (Text-fig. 13). 
At the point of entry into the penis sheath the duct becomes the epiphallus which 
lies coiled in the upper part of the sheath and which has a very different appearance 
in section (Text-fig. 14). The overall diameter is reduced but that of the lumen 
remains more or less unchanged. The internal epithelium is of irregularly cuboidal 
cells with central nuclei and no cilia. The nuclei are densely packed with chromatin 
and stain deeply in haematoxylin. The epithelial layer is surrounded by a 
layei of circular muscle and outside this is a layer of transverse muscle and connective 
tissue with bundles of longitudinal muscle fibres embedded in it. The transition 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 13 

from epiphalms to penis proper is gradual. Due to the mode of operation of the 
copulatory organ in Bulinus [Larambergue (1939) and Hubendick (19486)] the part 
that is proximal when the organ is at rest is distal when it is erected and the distal 
resting part is proximal during copulation. The following description is based on 
resting specimens. Proximally, the lumen increases gradually, the epithelial lining 
becomes regularly cuboidal and the central nuclei are less deeply staining. At 
about the maximum diameter of the penis the lumen is partially occluded by profound 
folding of the lining. The epithelium is of cuboidal cells with central nuclei 
containing sparse chromatin granules and beneath this is a layer of connective tissue 
with transverse muscle fibres and blood spaces. Around this is a layer of muscle 
and the outer layer is of mixed muscle fibres with blood spaces and connective 
tissue (Text-fig. 15). The distal tip of the penis, just before the point where it 
unites with the penis sheath, has a tri-radiate lumen surrounded by the same sort 
of epithelium as before and outside this only a thin layer of circular muscle (Text- 
fig. 16). The penis sheath has an outer covering of a thin, flattened epithelium 
within which is a layer of mixed muscle fibres followed by an innermost layer of 
almost pure circular muscle. The lining of the preputium is thrown into a number 
of folds with two muscular pillars predominating. Usually only one of these two 
pillars extends the whole length of the preputium, the other usually fades out 
before the junction with the penis sheath at the upper end. The position of the 
pillars within the preputium coincides with the point of attachment on the outside 
of two series of muscle fibres that connect the organ to the body wall of the head 
cavity. The epithelial lining of the preputium is fundamentally of columnar or 
cubical cells with more or less basal nuclei. In the most proximal parts of the organ 
a few ciliated cells are present. Distributed irregularly are gland cells, much larger 
than the other components of the epithelium, with basal nuclei often displaced to 
one side of the cell (Text-fig. 17). The cytoplasm of these gland cells may be either 
coarsely granular and eosinophilic or much more finely granular and staining deeply 
in haematoxylin. They are often clustered together in patches and may penetrate 
quite deeply into the muscular layers beneath. They are more common in the upper 
proximal parts of the organ but in one specimen examined they were confined almost 
exclusively to the epithelium covering one of the muscular pillars. Beneath the 
epithelium is a layer of circular muscle and outside this a thick layer of mixed muscles 
and connective tissue with blood spaces. 

Returning to the point of separation of the hermaphrodite duct, the female tract 
begins with a short duct with a narrow lumen lined by columnar glandular cells 
with darkly- staining basal nuclei. Between the free ends of these columnar cells 
are wedge cells with nuclei containing few chromatin granules. These wedge cells 
bear short cilia (Text-fig. 18). This short duct opens into the carrefour which has a 
wide lumen and a folded wall. The epithelial lining (Text-fig. 19) is of tall columnar 
cells and wedge cells. The glandular columnar forms have dark staining basal 
nuclei. There are patches of ciliated cells particularly in the folds of the wall. 
Opening into the carrefour is the duct from the albumen gland. This gland is made 
up of numbers of tubules with loose connective tissue between them. The lining of 
the tubules is quite characteristic in section ; it consists of roughly cuboidal cells 



i 4 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAU MEJ 

with basal granular nuclei and with numbers of large droplets in the cytoplasm. 
These droplets stain deeply in haematoxylin (Text-fig. 20). Distally to the carrefour 
the oviduct is lined with a columnar epithelium containing a few wedge cells and a 
few gland cells but without cilia (Text-fig. 21). The oviduct leads into the uterus 
which is elongate transversely in cross section. The epithelial lining is similar to 
that of the oviduct but there are more gland cells, some with acidophilic and some 
with basophilic granules in the cytoplasm. There are also a few ciliated cells (Text- 
fig. 22). For a considerable part of its course the uterus is surrounded by the muci- 
parous gland. Macroscopically this gland is smooth, colourless and rather 
translucent in appearance. In section it is seen to be made up of close-packed 
tubules with little intervening connective tissue. The tubules are lined with large, 
irregularly cubical cells with darkly staining basal nuclei. The cytoplasm is entirely 
without granules and is completely unstained by either haematoxylin or eosin 
(Text-figs. 23). The tubules open individually into the dorsal side of the uterus. 
Following immediately after the muciparous gland the uterus passes into the oothecal 
gland, distinguished macroscopically by its opaque white appearance in contrast to 
the translucent colourlessness of the previous gland. Internally the uterine wall is 
deeply folded in this region. The epithelium (Text-fig. 24) consists almost entirely 
of tall columnar glandular cells with basal nuclei. Both acidophils and basophils 
are present but the coarsely granular or eosinophilic type of cytoplasm predominates. 
A few patches of ciliated cells are present and the glandular cells may be several 
layers thick in places, penetrating deeply into the underlying connective tissue. The 
transition from the folded, glandular wall of the oothecal gland to the tubular, 
ciliated vagina is quite abrupt. The epithelial cells in this part are of a short 
columnar type with median to basal nuclei containing sparse chromatin granules. 
The cytoplasm of these ciliated cells is very finely granular and eosinophilic (Text-fig. 
25). The seminal receptacle duct which opens from the vagina is lined in its distal 
part (that nearest to the vagina) with a columnar epithelium with basal nuclei and 
finely granular acidophilic cytoplasm but without cilia. The rest of the duct lining 
is similar but the columnar cells are taller and ciliated. The distended sac of the 

Fig. 14. Transverse section of epiphallus of B. jousseaumei. 

Fig. 15. Part of transverse section of penis of B. jousseaumei. 

Fig. 16. Transverse section of tip of penis of B. jousseaumei. 

Fig. 17. Part of epithelial lining of the preputium of B. jousseaumei showing 
basophilic and eosinophilic gland cells. 

Fig. 18. Epithelial lining of proximal part of oviduct of B. jousseaumei. 

Fig. 19. Epithelial lining of carrefour of B. jousseaumei. 

Fig. 20. Epithelial lining of albumen gland tubule of B. jousseaumei. 

Fig. 21. Epithelial lining of distal part of oviduct of B. jousseaumei. 

Fig. 22. Epithelial lining of uterus of B. jousseaumei. 

Fig. 23. Epithelial lining of muciparous gland tubule of B. jousseaumei. 

Fig. 24. Epithelial lining of oothecal gland of B. jousseaumei. 

Fig. 25. Epithelial lining of vagina of B. jousseaumei, 



THE STRUCTURE AND TAXONOMY OF B.ULINUS JOUSSEAUMEI 15 




16 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

receptaculum is thin walled, a thin layer of connective tissue lying outside a tall 
columnar epithelium with basal nuclei and indistinct cell boundaries. 

The foregoing description is based on a study of sections of the genital organs of 
several adult snails in the 8-0-9-5 mm. shell length range. For comparison series 
of sections were cut through the reproductive organs of a juvenile snail (4-5 mm. 
shell length) and an " adolescent " specimen 5-7 mm. long. 

The gonad of the juvenile showed neither mature ova nor spermatozoa. Active 
cell division appeared to be in progress in the zone of the germinal epithelium near 
the bases of the acini but the state of fixation of the specimen made definite observa- 
tions difficult. No sections of the haemaphrodite duct were obtained. The male 
tract was not clearly defined until the level of the prostate gland. This organ was 
present but in section the tubules although defined were lined with an undifferentiated 
cuboidal epithelium with central nuclei. The vas deferens leading from the prostate 
was lined with a similar unciliated epithelium. The sheath of circular muscle 
present in the adult snail was represented by close-packed undifferentiated connective 
tissue cells. The duct remained more or less unchanged in this condition throughout 
its course. The epiphallus was again ensheathed in undifferentiated tissue and the 
epithelial lining was of more columnar cells with basal nuclei. The penis and penis 
sheath at this stage also showed no clear definition of muscle tissue and the separation 
between the muscle layers of the two parts was just becoming apparent. The folding 
of the inner wall of the penis was already in evidence. The preputium also was sur- 
rounded by undifferentiated muscular tissue and the lumen, already S-shaped owing to 
the development of the two main muscular pillars, was lined with a cuboidal epithelium 
with central nuclei. Of the female system at this stage little can be said. The 
uterus and vagina are present as tubes lined by a uniform epithelium. The albumen 
gland is entirely undeveloped, likewise the muciparous and oothecal glands. The 
receptaculum seminis and its duct are present but the receptacle sac is scarcely more 
than a slight dilatation of the duct. The only evidence of differentiation in the female 
tract in this specimen is that the nuclei of the epithelial cells in the regions that will 
become glandular stain more deeply in haematoxylin than do those of the other 
regions. 

The gonad of the " adolescent " snail (5-7 mm. shell length) showed clusters of 
mature spermatozoa in the acini with a few maturing oocytes in the upper parts. 
Some of the largest of these oocytes showed dividing nuclei and were, presumably, 
undergoing maturation division. Sections of the seminal vesicle and hermaphrodite 
duct were packed solidly with spermatozoa. The sperm duct after its separation 
from the common duct shows the same short, ciliated part followed by the long, 
glandular part with refractile eosinophil granules in the cytoplasm as is found in 
the adult. The prostate gland also is identical histologically to the adult. Through- 
out its course the vas deferens corresponds in histological detail to the form already 
described in the adult. The transition to the epiphallus is similar as are the structures 
of the penisand penis sheath. The preputium, however, is, in its distal part, similar 
to that in the juvenile in that the lumen is S-shaped and only two muscular pillars 
are present. More proximally additional folds in the wall do occur. The epithelium 
lining the lumen is more ciliated than in the adult and few gland cells have been 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 17 

observed. The female tract at this stage is far less well developed. The albumen 
gland is represented by a fairly compact mass of tissue of undoubtedly glandular 
nature but scarcely organized into tubules as in the adult. The few tubules which 
are present are not lined with the characteristic glandular epithelium of the adult 
but with cells which probably later develop into this form. The carrefour is present 
as a dilatation of the oviduct but its epithelial lining is only differentiated into 
glandular areas in parts, some cilia are also present. The oviduct is similar histo- 
logically to that in the adult and it in turn passes into the ciliated part of the uterus. 
Neither the muciparous nor the oothecal glands is represented by more than a slight 
thickening of the uterine wall. The receptaculum seminis is well developed and 
contains spermatozoa and the vagina is strongly ciliated. 

These histological observations support the opinion already formed from gross 
anatomical studies that the male genital tract develops slightly earlier than the 
female system. Morton (1954) has suggested that a protandrous sexual cycle is the 
primitive condition in gastropods and that simultaneous hermaphroditism has been 
later developed in the higher pulmonates and opisthobranchs. Larambergue (1939) 
states that in Bulinus contortus there is no protandry since spermatozoa and ova are 
produced simultaneously throughout life even though spermatozoa do appear first. 
It seems probable that the earlier development of the male tract is a relic of the 
primitive protandrous condition which has become almost completely obscured, 
particularly where the length of the breeding season is limited by adverse 
environmental conditions. 

To complete this histological study of the genital organs of B. jousseaumei a few 
comparisons with similar studies on related forms should be considered. It has 
already been said that there are no differences between the fundamental structure of 
the gonad in this species and in Biomphalaria boissyi and Helisoma trivolvis as 
described by Abdel-Malek (loc. cit.) and it is probable that this structure is fairly 
uniform throughout the Planorbidae. The hermaphrodite duct corresponds to the 
description by Larambergue (loc. cit.) for that in B. contortus, also the point of 
separation of the male and female tracts. The sperm duct differs from that in B. 
contortus in that no ciliation has been observed in its lumen apart from the very 
short part immediately after its separation from the common duct. Abdel-Malek 
mentions no ciliation of the sperm duct in B. boissyi and only near the prostate in 
H. trivolois. In both of these species he mentions the refractile, eosinophilic, 
cytoplasmic granules but also records the presence of scattered basophils not seen in 
B. jousseaumei. The prostate corresponds well with that described by Larambergue 
for B. contortus and this author stresses the fact that the gland is not traversed by 
the male duct but that its tubules open into a central chamber into which the sperm 
duct opens at one side and from which the vas deferens leaves at the other. The 
vas deferens in B. jousseaumei differs from that in B. contortus in that it is ciliated 
throughout its length while Larambergue reports that in the second species the 
epithelium is unciliated and of a mucous-secreting type. In both of the species 
described by Abdel-Malek the vas deferens is ciliated throughout its length although, 
due to the different form of the prostate in these species, there is only a gradual 
transition from the form of the sperm duct to that of the vas deferens instead of a 



18 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

clear-cut demarcation of the two as in B. jousseaumei. The epiphallus which is so 
well differentiated from the vas deferens in the present species was also noted 
(although not under this name) by Larambergue to be histologically different. 
Since the structure of the copulatory apparatus in the species described by Abdel- 
Malek differs so markedly from that in the Bulinids it is not possible to draw com- 
parisons between the histology of the two but it is interesting to note that in his 
species the seminal canal within the penis (and therefore that part of the male duct 
actually inside the penis sheath) is ciliated and that the epithelial lining does not 
differ from that of the vas deferens. The histological structure of the penis, penis 
sheath and preputium in the present species show no significant differences from 
those described by Hubendick (19486) and Larambergue. Larambergue does not 
describe the histology of the female genital tract of B. truncatus apart from mention- 
ing the ciliation of the receptaculum seminis duct. The epithelial linings of the 
first part of the oviduct and of the carrefour are similar to those of B. boissyi and 
H. trivolvis although not so heavily ciliated. Beyond the carrefour the epithelium 
of the oviduct in the present species almost entirely lacks cilia while these are present 
in Biomphalaria. The epithelium of the uterus is similar in the two species, and the 
muciparous gland merges gradually with the uterus rather than abruptly as in 
Helisoma. The remainder of the female tract is very similar to Biomphalaria except 
that the part of the receptaculum seminis duct nearest to the vagina is unciliated, 
a condition similar to that in Helisoma. 



RELATIONSHIPS OF B. JOUSSEAUMEI 

In considering the affinities of B. jousseaumei it is necessary to review the species 
of the sub-genus Physopsis known from West Africa. For this purpose it is often 
necessary to refer to the medical literature as it is largely in works on the epidemiology 
and transmission of human schistosomiasis that records of intermediate hosts are 
to be found. The only species of Physopsis actually described from West Africa is 
P. globosa (Morelet, 1866) collected in Angola. Of the other twenty or so species, 
all were described originally from East or South Africa. The question of the 
identity of P. globosa with P. africana Krauss, 1848, will not be dealt with here 
since opinions are still divided on this point and much detailed work must be carried 
out before a solution of the problem can be reached. It must, however, be pointed 
out that many authors now treat P. globosa as an absolute synonym of P. africana 
others consider it to be a variety or sub-species of the latter while still others 
recognize it as a separate species. In the following account it will be pointed out 
(where it is known) which of these three courses was adopted by the workers 
concerned. Since a chronological account of these records would undoubtedly prove 
confusing, they are presented in a geographical sequence from north to south. 

McCullough & Duke (1954) were the first to record Bulinus africanus from the 
Gambia and they noted that this was probably the northernmost record for the 
species in West Africa. They were following the classification of Amberson & 
Schwarz (1953) who treated all species of Physopsis as forms of B. africanus and there 
is no doubt that their record refers to B. jousseaumei. I later recorded (Wright, 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 19 

1956) a form of B. (P.) globosus from one localty in Upper River Division, Gambia, 
as well as B. jousseaumei from the Casamance Province of Senegal, the next territory 
to the south of the Gambia. No published records of Physopsis from Portuguese 
Guinea have been found, and Pinto (1949) in a recent survey of vesicle schistosomiasis 
in that territory found only Bulinus forskali and B. dautzenbergi. Vogel (1932) in 
a similar survey in French Guinea and Liberia found (and illustrated) Physopsis 
globosa. More recently in a rather confusing account of schistosomiasis in French 
West Africa, Le Gall (1944) mentions as a probable vector of urinary schistosomiasis 
in French Guinea " Physopsis ovoides and boissyi " from Kissidougon. This 
presumably refers to P. ovoideus and Biomphalaria boissyi. The next territory 
southwards from the French Guinea coast, Sierra Leone, is perhaps one of the best 
documented areas in tropical Africa with respect to the epidemiology of 
schistosomiasis. It has been the subject of three major surveys. Blacklock (1924), 
Blacklock & Thompson (1924), Blacklock (1925), Gordon, Davey & Peaston (1934) 
and Gerber (1952). Connolly (1928) published an account of the freshwater 
molluscan material collected by Blacklock and considered the Physopsis to be 
P. globosa with strong affinities to P. didieri. He mentioned the presence of a well- 
marked spiral sculpture on the material from Sierra Leone, a feature which he had 
not observed on P. globosa from Angola and Mozambique. Connolly also identified 
material submitted to him by Gordon and his co-workers and identified the Physopsis 
as P. globosa. Gerber submitted his material to Dr. W. J. Rees who identified one 
fully-grown and three smaller specimens out of a batch of 204 shells as Bulinus 
globosus and referred the remaining 200 to B. africanus. The same sample of shells 
was shown to Berry who pronounced them all to be Physopsis africana. To the 
south of Sierra Leone is Liberia and here Vogel (loc. cit.) records Physopsis globosa 
and Veatch (1946) mentions Physopsis africana var. globosa as the intermediate 
host of Schistosoma haematobium in the Western Province. No records of identified 
Physopsis from the Ivory Coast have been seen but Ingram (1924) incriminated 
P. globosa as the possible vector of urinary schistosomiasis in the Gold Coast ; more 
recently Edwards & McCullough (1954) have demonstrated that the parasite is 
carried by P. africana but they also mention that they consider P. globosa to be 
either a race or an absolute synonym of this species. The only further record of an 
identified Physopsis between the Gold Coast and the Belgian Congo is that of P. 
globosa collected at Kano in Northern Nigeria (Gordon, 1932). In the excellent 
work of Pilsbry & Bequaert (1927) a variety of P. africana is recorded from a number 
of localities in the Belgian Congo, also P. africana globosa. These authors refer the 
other recorded species of Physopsis (apart from P. tanganyicae von Martens) from the 
Belgian Congo either to P. africana var. or to P. africana globosa. Finally, the 
territory to the south of the Belgian Congo, Angola, is the type locality for P. globosa 
Morelet. 

Examination of material from French Guinea, Sierra Leone, Liberia, Belgian 
Congo and Angola has shown that without doubt the same species of Physopsis is 
present in all these territories and that B. jousseaumei is a form of that species. 
The table below gives the mean dimensions in millimetres of the populations examined 
from the territories mentioned above ; 



2o THE STRUCTURE AND TAXONOMY 


OF BULINUS JOU 


SSEA UMEI 


Territory 
and number 
of specimens 


Shell 
length 


Shell 
max. 
diam. 


Aperture 
length 


Aperture 
width 


Shell length 
Aperture width 


Shell length 
Aperture length 


Gambia . 
283 


7-84 
±1-52 


6- 00 

±1*12 


. 6-65 . 
• ±1-03 • 


3-37 
±o-66 


2-31 
±0-14 


i- 17 
±0-08 


Casamance 
20 . 


8-69 
±0-96 


6-32 
±0-76 


7-21 . 
. ±0-70 . 


3- 60 
±o-33 


2-41 
±0-13 


i- 20 

±0-04 


French Guinea 
20 . 


io-33 

±o-75 


7-01 
±0-42 


. 8- 3 i . 
• ±0-51 . 


4.04 
±0-20 


2-55 
±o-n 


1-24 
±0-05 


Sierra Leone . 
100 


10-07 
±2-06 


. 7.18 

±i-75 


. 8-43 . 
• ±i-43 • 


3-97 
±0-93 


2-55 
±0-19 


1 -19 
±0-08 


Liberia . 
6 . 


10-56 

±2-81 


- 7-38 
±1-63 


8-8 . 
. ±1-98 . 


4-28 
±1-06 


2-47 
±0-15 


1. 19 

±0-07 


Belgian Congo 
100 


n-34 
±1-64 


■ 8-50 

±1-22 


. 9-5i • 
• ±1-52 • 


4'54 
±0-63 


2-48 
±o-n 


1-19 
±0-065 


Angola (Type series 
26 . 


) 13-43 
±2-19 


- 9-5 1 
±1-63 


9- 60 . 
. ±1-48 . 


5-13 
±0-85 


2-62 

±0-15 


1 -40 
±0-085 



Text-fig. 26 shows the means for the ratio Shell length /Aperture length plotted 
against the mean lengths for each of these samples. Text-figs. 27-30 show the 
ratios shell length /aperture length and the frequency distributions of shell length in 
the samples from Sierra Leone and the Belgian Congo. 

Reference to the table shows a gradual decrease in the dimensions of the shell from 
south to north. The composition of the samples is not wholly comparable, those 
from the Gambia, Sierra Leone and Belgian Congo are more or less random popula- 
tion samples with a proportion of juveniles and those from Casamance, French 
Guinea and the type series from Angola contain mostly adult specimens. The small 
sample from Liberia consisted of four adults and two juveniles, hence the rather 
large standard deviations. The means of the type series differ from those given by 
Mozeley (1939) because his figures were based on ten specimens of the series only. 
In addition to the discrepancies in sampling certain differences due to ecological 
conditions also occur. The most obvious of these is the " still water " effect. 
Schwetz (1954) has described the effect of changed environment on shell form in 
several planorbid snails and the exsertion of the spire in forms which develop in 
static water compared to those in gently moving streams is well known. This 
accounts largely for the difference in the mean values of the ratio shell length/ 
aperture length in the type series and in the Belgian Congo sample. The latter came 
from a pool in a recently dried stream bed while Morelet (1S68) notes that the Angolan 
material was collected in a lake. Similarly this probably accounts for some of the 
differences between Gerber's and Blacklock's specimens from Sierra Leone. Gerber's 
material from the still waters of a rice swamp has a more exserted spire than 
jBlacklock's from a stream, 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 21 



M 












A 


1- J 














L ML 








FC 






1.1 


26 


c 


c 


SL L 


BC 





2 3. 4 S « ' ■ » 10 II |2 |] 14 || u | 7 

SHELL LENGTH IN MILLIMETRES. 



27 



t/ML 
1.2 



28 




L ML 
1.2 



30 



7 8 » 10 II 12 II 

SHELL LENGTH IN MILLIMETRES 



Fig. 26. Graph of mean ratio shell length /aperture length against mean shell length 
for populations of B.jousseaumei from Gambia (G), and Casamance (C) and B. globosus 
from French Guinea (FG), Sierra Leone (SL), Liberia (L), Belgian Congo (BC) and the 
type series from Angola (A) . 

Fig. 27. Size-frequency histogram for a population of B. globosus from Belgian Congo. 

Fig. 28. Graph of ratio shell length /aperture length against shell length for the same 
population. 

Fig. 29. Size-frequency histogram for sample of B. globosus from Sierra Leone. 

Fig. 30. Graph of ratio shell length /aperture length against shell length for the same 
population. 



22 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

The general form of the shell is similar throughout the whole range under considera- 
tion (Plate I). In the same population shells may be seen with the columella 
margin fused entirely to the body whorl and others with a space between the two. 
This was stressed by Connolly (1934) as being the only constant conchological 
difference between Physopsis africana and P. globosa. There is also a range of 
variation in the angle at which the outer lip meets the body whorl at the top of the 
aperture, a character used by Pilsbry & Bequaert (1927) for the separation of the 
same two species. The sculpture pattern described earlier for the Gambian material 
is found also throughout the range, being most marked in the specimens from 
Sierra Leone (Plate 2,) and, as noted by Connolly (1928) practically absent from 
the Angolan material. This is of interest since Mandahl-Barth (1954) erected a sub- 
species Bulinus globosus ugandae which he separated from the nominate sub-species 
because it completely lacked the spiral sculpture of the typical form. In passing 
it seems appropriate to note that amongst the shells collected in Northern Rhodesia 
by Buckley (1946) were a number of specimens labelled by Connolly as juvenile 
Bulinus natalensis but which were also referrable to B. hemprichii depressus Haas, 
1936. These specimens were indistinguishable from many of the juvenile Phy- 
sopsis globosa seen during this work. Unfortunately the type specimens of Haas' 
subspecies were destroyed or lost during the war but from a study of the photo- 
graphs of these specimens there can be little doubt that the sub-species should be 
placed in the synonymy of Bulinus (Physopsis) globosus. 

It can be seen from Text-fig. 30 that the pattern of the graph of shell length/ 
aperture length against shell length for the sample from Sierra Leone is similar to 
that for the Gambian population (Text-fig. 3) . For the lower values of shell length 
the graph is almost parallel to the horizontal axis and the gradual upward slope does 
not begin until a shell length of about 8-o mm. is reached. It has already been 
shown that this change is brought about by the onset of the adult phase and it should 
be noted that it occurs at a greater shell length in the Sierra Leone population than 



Figs. 31-35. Stages in the development of the accessory genital glands and 
male copulatory organ of B. jousseaumei from the Gambia. 

Fig. 31 at 4- 5 mm. shell length. 
Fig. 32 at 5-2 mm. shell length. 
Fig. 33 at 6- 3 mm. shell length. 
Fig. 34 at 7 • 5 mm. shell length. 
Fig. 35 at 8*8 mm. shell length. 

Figs. 36-38. Stages in the development of the accessory genital glands and 
male copulatory organ of B. globosus from Sierra Leone. 

Fig. 36 at 6- 1 mm. shell length. 
Fig. 37 at 8-5 mm. shell length. 
Fig. 38 at 9-2 mm. shell length. 

Figs. 39-41- Similar stages in B. globosus from Angola. 

Fig. 39 at 8-9 mm. shell length. 
Fig. 50 at 9*5 mm. shell length. 
Fig. 41 at 9-75 mm. shell length. 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 23 







24 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

in that from the Gambia. That the same characteristic is present in the correspond- 
ing graph for the Belgian Congo material is almost certain ; the distribution of 
individuals in the sample is, however, such that the feature is not well marked al- 
though it seems probable that it occurs at an even greater size than in the Sierra 
Leone population. 

Although no anatomical differences in structure have been found in the limited 
material examined (Sierra Leone, Belgian Congo and Angola) there is a variation in 
degree of development of the genital organs at corresponding shell sizes. Text-figs. 
31-35 show stages in the development of the accessory genital glands and male 
copulatory organ at various shell lengths for B. jousseaumei in the Gambia. Text- 
figs. 36-38 are of three stages in B. globosus from Sierra Leone and Text-figs. 39-41 
for material from Angola. The degree of development of the Gambian specimen 
of 5-2 mm. shell length (Text-fig. 32) compares well with that of 8-5 mm. from 
Sierra Leone (Text-fig. 37) and 9-5 mm. from Angola (Text-fig. 40). The protandrous 
development of the male copulatory organ and prostate is even better marked in 
these last two groups of specimens than it is in B. jousseaumei. 

Of possible significance, but at present insufficiently investigated, is the increased 
pigmentation of the mantle in the more southern forms. Specimens from Sierra 
Leone and the Belgian Congo have the mantle heavily blotched with black in contrast 
to the more diffuse spotting of that in B. jousseaumei. Three of the four specimens 
dissected from Angola had almost the entire mantle black with one or two lighter 
patches. 

Differences in the radula do not seem significant ; one or two more laterals in 
each transverse half row may be found in the larger forms but this is to be expected. 

DISCUSSION 

Evidence has been presented to show that Bulinus jousseaumei from the Sene- 
gambian region is closely related to the other species of the sub-genus Physopsis 
from neighbouring West African territories and that these in turn are related to the 
typical Bulinus globosus. In spite of a number of distributional gaps, probably due 
to an absence of collectors rather than of snails, it seems clear that there is a well- 
marked cline grading from the typical globosus form in the south to the small 
jousseaumei at the extreme northern limit of the range. Not only is this cline 
represented by a gradation in size but also by a gradation in the degree of protandric 
development, possibly also by differences in the intensity of mantle markings. 
The change does not become really well marked until (moving northward) Sierra 
Leone is reached. It seems very probable that the cline is correlated with the length 
of the rainy season. In the Senegambian region there is a single wet season of about 
four months duration while to the south the season is prolonged and may be 
duplicated. The short single wet season will limit the time during which the 
streams and bolons are suitable for the development of snails, hence the telescoping 
of the sexual phases and the reaching of sexual maturity at a smaller shell size. In 
regions of more continuous rainfall the need for rapid development is less, resulting 
in the more marked protandrous development (a primitive character) and the later 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 25 

onset of sexual maturity, the latter being of course closely connected with the larger 
shell size. 

Although the use of a trinomial system of nomenclature is of doubtful value unless 
it is well documented it seems justifiable to retain the name jousseaumei as a sub- 
species in this case, for the Senegambian form. It differs from the typical form of 
globosus in its considerably smaller size and its apparently more contracted life 
cycle. It is at present geographically isolated from the typical form by the distribu- 
tional gap in Portuguese Guinea but subsequent work may well show that this is 
not actually so. The name Bulinus (Physopsis) globosus jousseaumei (Dautzenberg) 
is therefore proposed. 

If B. jousseaumei is a sub-species of B. globosus then the problem of its affinities 
with the strongly umbilicate form of B. globosus described from the Gambia (Wright, 
1956) arises since two geographical races of the same species are not to be expected 
in the same geographical region. In this form the columellar margin of the aperture 
is greatly developed and not reflected, giving rise to a wide umbilicus with a slight 
keel around its opening. The columellar truncation is also thus suppressed leaving 
only a thickened line on the inner surface of the columella. There appears to be no 
record in the literature that the type series of B. globosus includes several specimens 
which show this character in a very limited degree. Material in a collection made in 
Northern Rhodesia by Dr. P. Le Roux contains a number of specimens which show 
this character even further developed. Intermediate forms between the Rhodesian 
and Gambian specimens have been seen in Blacklock's collection from Sierra Leone. 
In these the umbilicus is well developed but not quite so wide as in the Gambian 
material. Although the evidence is incomplete it appears that this variety of 
B. globosus also shows a clinal distribution parallel to that described for the typical 
form and B. jousseaumei. In this instance the main character in which gradation 
has been observed is in the degree of overgrowth of the columellar margin with con- 
sequent suppression of the columellar truncation and increase in the size of the 
umbilicus. Insufficient spirit material has been available for a study of associated 
anatomical variation. The only locality from which this umbilicate form was 
obtained in the Gambia was at Badja Kunda, Upper River Division where the 
ecological conditions differed slightly from those in the typical B. jousseaumei 
habitats. A single specimen of B. jousseaumei showing this character slightly 
developed was obtained with normal specimens at Sudowol bridge over the Simoto 
bolon, Upper River Division. The evidence suggests that this is possibly a recessive 
genetic character of the normal form which, when it occurs as a pure homozygous 
population is better adapted to slightly different habitats and appears to behave as 
a separate species. 

SUMMARY 

1. A brief biometrical study of the shell of B. jousseaumei is made. 

2. The histology and course of development of the genital tract of this snail is 
described and compared with related planorbids. 

3. The records of Physopsis spp. from West Africa are briefly surveyed. The 
possibly fallacious premises on which Bulinus globosus ugandae Mandahl-Barth 



26 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

was described are mentioned and Bulinus hemprichii depressus Haas is referred to 
the synonymy of B. globosus. 

4. The relationship of B. jousseaumei to B. globosus is discussed and the former 
is reduced to a sub-species of the latter as B. g. jousseaumei, the northernmost 
representative of a cline of the typical form. 

5. The relationship of B. g. jousseaumei to the umbilicate form of B. globosus is 
discussed and this form is related through a graded series to the typical form. 

ACKNOWLEDGMENTS 

In addition to acknowledgments made in the text I am greatly indebted to Dr. 
T. P. Eddy, Director of Medical Services, Sierra Leone, and to Dr. J. Schwetz of 
Brussels, both of whom have sent to me useful material from Sierra Leone and the 
Belgian Congo respectively ; also to Dr. John Morton who has been kind enough to 
read the manuscript and make a number of useful suggestions. My thanks are also 
due to the Director of the Institut Royal des Sciences Naturelles de Belgique both 
for the photograph of the type specimens of Isidora jousseaumei and for permission 
to publish this photograph. 

REFERENCES 
Abdel-Malek, E. T. 1954a. Morphological studies on the family Planorbidae (Mollusca : 
Pulmonata). I. Genital organs of Helisoma trivolvis (Say) (Subfamily Helisomatinae F. C. 
Baker, 1945). Trans. American Microscopical Soc. 73 (2) : 103-123. 

1954&. Morphological studies on the family Planorbidae (Mollusca : Pulmonata). II. 

The genital organs of Biomphalaria boissyi (Subfamily Planorbinae, H. A. Pilsbry 1934), 
Ibid. 73 (3) : 285-296. 

Amberson, J. M. & Schwarz, E. 1953. On African Schistosomiasis. Trans. Roy. Soc. Trop. 

Med. Hyg. 47 (6) : 451-502. 
Blacklock, D. B. 1924. Report of an investigation into the prevalence and transmission 

of human schistosomiasis in Sierra Leone. Sierra Leone Ann. Med. San. Rep. for 1923 : 

80-87. 

1925. Endemic goitre and schistosomiasis in Sierra Leone. Trans. Roy. Soc. Trop. Med. 

Hyg. 18 (8) : 3 95"427- 

— ■ — ■ & Thompson, M. G. 1924. Human schistosomiasis due to 5. haematobium in Sierra 

Leone. Ann. Trop. Med. &> Parasit. 18 (2) : 211-234. 
Buckley, J. J. C. 1946. A helminthological survey in Northern Rhodesia. /. Helminthology, 

21 (4) : 111-174. 
Connolly, M. 1928. The non-marine mollusca of Sierra Leone. Ann. Mag. Nat. Hist. 

Series 10, 1 (4) : 529-551. 

1934- On the planorbid hosts of bilharziasis in South and West Africa. Ann. Trop. Med. 

& Parasit. 28 (3) : 439-443. 

Dautzenberg, P. H. 1890. Recoltes malacologiques de M. le capitaine Em. Dorr, dans le 
Haut-Senegal et le Soudan Francais de 1 886-1889. Mem. Soc. Zool. France, 3 : 132-134. 

Deschiens, R. 1951. Le probleme sanitaire des bilharzioses dans les territoires de l'Union 
Francaise.JBw//. Soc. Path. Exot. 44 : 631-638. 

Dupuis & Putzeys. 1923. Deuxieme note concernant le Faune Malacologique Africaine. 
Ann. Soc. Zool. et Malac. Belgique, 53 : 69-79. 

Edwards, E. E. & McCullough, F. S. 1954. Studies on the life cycles of Schistosoma haema- 
tobium and 5. mansoni in the Gold Coast. Ann. Trop. Med. and Parasit. 47 : 164-177. 

Gerber, J. H. 1952. Bilharzia in Boajibu. Part 1. /. Trop. Med. 6- Hyg. 55 (3) : 52-58. 

1952. Bilharzia in Boajibu. Part 2. Ibid. 55 (4) : 79-93. 



THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 27 

Gordon, R. M. 1932. The molluscan host of Schistosoma haematobium in Northern Nigeria. 
Ann. Trop. Med. d>* Parasit. 26 : 117-118. 

Davey, T. H. & Peaston, H. 1934. The transmission of human bilharziasis in Sierra 

Leone, with an account of the life-cycle of the schistosomes concerned, 5. mansoni and S. 
haematobium. Ibid. 28 (3) : 323-418. 

Haas, F. 1936. Binnen-Mollusken aus Inner Afrika. Abh. Senckenberg. Naturf. Ges. 431 : 

1-156. Frankfurt A.M. 
Harding, J. P. 1949. The use of probability paper for the graphical analysis of polymodal 

frequency distributions. J.M.B.A. 28 : 141-153. 
Hubendick, B. 1948a. The Anatomy of Bulinus, with a discussion of the term prostate and 

its sense in the Basomatophora. Proc. Malac. Soc. Lond. 27 (5) : 186-196. 

19486. Studies on Bulinus. Ark. for zoologi, 40 A (16) : 1-63. 

1951. Recent Lymnaeidae. Kungl. Svenska Vetenskap. Hand. Fjarde Serien, 3 (1) : 

1-223. 

Ingram, A. 1924. Note on a possible intermediate host of Schistosoma haematobium in the 

Gold Coast. Ann. Trop. Med. &> Parasit. 18 (3) : 265-266. 
Larambergue, M. de. 1939. Etude de l'autofecondation chez les gasteropodes pulmones. 

Recherches sur l'aphallie et la fecondation chez Bulinus (Isidora) contortus Michaud. Bull. 

Biol. Paris, 73 : 1-231. 
Le Gall, R. 1944. Les bilharzioses en Afrique Occidentale Francaise au Togo et a Madagascar 

de 1939a 1941. Bull. Off. Int. d'Hyg. Pub. 36 : 1 16-126. 
Mandahl-Barth, G. 1954. The freshwater mollusks of Uganda and Adjacent territories. 

Ann. Mus. Roy. Congo Beige Tervuren, Serie in 8°, Sciences zoologiques 32. 
McCullough, F. S. & Duke, B. O. L. 1954. Schistosomiasis in the Gambia. 1. Observa- 
tions on the potential snail vectors of Schistosoma haematobium and 5. mansoni. Ann. 

Trop. Med. & Parasit. 48 (3) : 277-286. 
Meeuse, A. D J. 1950. Rapid methods for obtaining permanent mounts of radulae. Basteria, 

14 (2 & 3) : 28-43. 
Mo relet, A. 1866. Coquilles nouvelles recueilles par le Dr. Fr. Welwitsch dans 1' Afrique 

equatoriale et particulierement dans les provinces portugaises d'Angola et de Benguela. 

/. de Conchy liologie, 14 : 153-163. 

1868. Mollusques terrestres et fluviatiles. Voyage du Dr. Friederick Welwitsch, Paris. 

Morton, J. E. 1954. The pelagic mollusca of the Benguela Current. With an account of 

the reproductive system and sexual succession of Limacina bulimoides. Discovery Reports, 

27 : 163-200. 
Mozeley, A. 1939. The freshwater mollusca of the Tanganyika Territory and Zanzibar 

Protectorate and their relation to human schistosomiasis. Trans. Roy. Soc. Edinburgh, 59 (3) : 

687-744. 
Peters, B. G. 1938. Biometrical observations on shells of Limnaea Species. /. Helminth. 

16 (4) : 181-212. 
Pilsbry, H. A. & Bequaert, J. 1927. The aquatic molluscs of the Belgian Congo. Bull. 

Amer. Mus. Nat. Hist. 53 (2) : 69-602. 
Pinto, A. R. 1949. Os primeiras dados sobre a existencia da Schistosomiase vesical na 

Guine Portuguese e importancia da contagem de ovas do parasito no sedimento urinario. 

Anais. Inst. Med. Trop. Lisboa 6 : 75-114. 
Schwetz, J. 1954. L'influence du milieu sur la taille et la forme du meme Planorbe ou du 

meme Bulinus. Ann. Soc. Roy. Zool. Beige, 85 (1) : 23-34. 
Smithers, S. R. 1956. On the ecology of schistosome vectors in the Gambia with evidence of 

their role in transmission. Trans. Roy. Soc. Trop. Med. & Hyg. 50 (4) : 354-365. 
Veatch, E. P. 1946. Human trypanosomiasis in Liberia 1941-44. Supplement to Amer. J. 

Trop. Med. 26 (5) : 1-56. 
Vogel, H. 1932. Beitrage zur epidemiologic der schistosomiasis in Liberia und Franzosisch- 

Guinea. Arch. Schiffs-u. Tropenhyg. 36 (3) : 108-135. 



28 THE STRUCTURE AND TAXONOMY OF BULINUS JOUSSEAUMEI 

Watson, H. (In Connolly, M.). 1925. The non-marine mollusca of Portuguese East Africa. 

Trans. Roy. Soc. S. Africa, 12 (3) : 105-220. 
Wright, C. A. 1956. The anatomy of six species of the Molluscan genus Bulinus (Planorbidae) 
from Senegambia. Proc. Malac. Soc. Land. In press. 



ABBREVIATIONS USED IN FIGURES 



Figs. 5 &6. 



Fig. 7. 



Fig. 35. 



BUC 


= buccal ganglion. 


CER 


= cerebral ganglion. 


OT 


= otocyst. 


OTN 


= otocyst nerve. 


PED 


= pedal ganglion. 


PENN 


= penial nerve. 


PL 


= pleural ganglion. 


VIS 


= visceral ganglion. 


CAE 


= caecum. 


CRO 


= crop. 


GIZ 


= gizzard. 


INT 


= intestine. 


STO 


= stomach. 


ALBG 


= albumen gland. 


HD 


= hermaphrodite duct. 


MUCG 


= muciparous gland. 


OOTG 


= oothecal gland. 


PR 


= preputium. 


PROS 


= prostate. 


PS 


= penis sheath. 


RS 


= receptaculum seminis 


SD 


= sperm duct. 


VAG 


= vagina. 


VAS DEF 


' = vas deferens. 




PLATE i 



Top row : Bulinus (Physopsis) globosus jousseaumei from Upper River Division, Gambia 

(X 2.) 

2nd row : B. (P.) globosus jousseaumei from Casamance Province, Senegal. ( x 2.) 

3rd row : B. (P.) globosus from French Guinea. ( x 2.) 

4th row : B. (P.) globosus from Kailahun District, Sierra Leone. ( x 2.) 

5th row : B. (P.) globosus from Bolahun, Liberia. ( x 2.) 

6th row : B. (P.) globosus from Kongola, Belgian Congo, (x 2.) 

7th row : B. (P.) globosus from Angola, specimens from type series. ( x 2.) 



Bull. B.M. (N.H.) Zool. 5, 1. PLATE 1. 



$v$$i%$<N> 




*$& #t)$^ 




PLATE 



Top row 
2nd row 
3rd row 
4th row 
5 th row 

(X 2.) 

Bottom : 



Figured type specimens of Isidora jousseaumei Dautzenberg. (x 1-5.) 
Umbilicate form of Bulinus (Physopsis) globosus from the Gambia. ( x 2.) 
Umbilicate form of B. (P.) globosus from Sierra Leone. ( x 2.) 
Umbilicate form of B. (P.) globosus from Northern Rhodesia. ( X 2.) 
Umbilicate form of B. (P.) globosus from Angola, specimens from type series. 

High -power view of micro-sculpture on shell of B. (P.) globosus from Sierra Leone 



Bvdl.B.M. (N.H.) Zool. 5, i 



PLATE 2. 











2 8 FEB\957 



&l *§ 



( 2 2MARW57] 

\^ 

ON SPELAEOGRIPHUS, 

A NEW CAVERNICOLOUS 

CRUSTACEAN FROM 

SOUTH AFRICA 



ISABELLA GORDON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 2 

LONDON : 1957 



ON SPELAEOGRIPHUS , A NEW 
CAVERNICOLOUS CRUSTACEAN FROM 
SOUTH AFRICA 



BY 

ISABELLA GORDON 



Pp. 29-47 ; 26 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 2 

LONDON : 1957 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

This paper is Vol. 5, No. 2 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March 1957 Price Six Shillings 



ON SPELAEOGRIPHUS, A NEW CAVERNICOLOUS 
CRUSTACEAN FROM SOUTH AFRICA 

By ISABELLA GORDON, D.Sc, Ph.D. 

SYNOPSIS 

A new cavernicolous Malacostracan from a pool in a cave on Table Mountain, Spelaeogriphus 
lepidops n.g. and sp., is described and figured. The affinities of the genus are discussed. In some 
respects it resembles Monodella (Thermosbaenacea), in others it approaches the Anaspidacea 
(Syncarida), or the Tanaidacea (Peracarida), but it is not referable to any of these Orders. 
Specimens received later included an ovigerous female carrying ten to twelve large ova in a 
characteristic Peracaridan brood-pouch composed of five pairs of oostegites. The genus, therefore, 
belongs to the Division Peracarida and, as it is not referable to any of the existing Orders of 
that Division, is placed in a new Order Spelaeogriphacea and a new family Spelaeogriphidae 
each with the characters of the genus. Nothing is known of the internal anatomy or of the 
embryology. 

INTRODUCTION 

Recently members of the South African Spelaeological Association obtained some 
specimens of a small blind Malacostracan from a pool at a depth of no ft. in a cave 
on Table Mountain, South Africa. The animals were said to " swim swiftly with 
rapid undulations of the body ". The temperature of the water in which they 
lived was 50 F. (February, 1956). 

The specimens were submitted to Dr. K. H. Barnard who found that they 
represented a new genus and species of Crustacea Malacostraca which at first sight 
seemed referable to the Division Syncarida but on closer examination seemed rather 
to belong to the Peracarida — its " affinities seem to be with the Tanaidacea, especially 
Apseudes " to quote from Dr. Barnard's letter. In April 1956 Barnard sent to the 
British Museum six of the specimens, together with notes and sketches, and 
suggested that I might like to describe this interesting new species and discuss its 
affinities in more detail. I wish to express my thanks to Dr. Barnard for presenting 
these specimens to the Museum and also for the privilege of studying them. 

The specimens were sent to London in two small vials. One vial contained two 
almost-perfect specimens which are quite opaque and much better preserved than 
the other four ; these have been selected as the holotype, a male measuring 7-2 mm. 
in length (from tip of rostrum to posterior margin of telson) and the allotype, a 
female measuring 5-6 mm., respectively. The four paratypes are very delicate, 
almost transparent, and more or less imperfect as to their appendages ; they range 
from 4-9 to 6-8 mm. in length and comprise two females and two males (one rather 
immature). The holotype and allotype have been handled with care and, for the 

ZOOL. 5, 2. 2§ 



32 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 

necessary dissection, the more incomplete paratypes have been used. Some 
specimens have been retained for the South African Museum collection and according 
to Barnard the largest measures 7-5 mm. in length. 

The name suggested by Barnard for this new genus was most appropriate but, 
unfortunately, it is now preoccupied by Spelaeocaris Matjasic (1956, p. 65) a new 
genus of the family Atyidae. I therefore propose the name Spelaeogriphus from 
" griphos " meaning something complicated, a puzzle or riddle, not " gryphos " 
meaning a griffin (Jaeger, 1955). 

Spelaeogriphus n. g. 

Diagnosis. Body elongated, subcylindrical (somewhat depressed). Carapace 
short, deep, coalescing dorsally with first thoracic somite and overhanging on each 
side to largely conceal the mouthparts and to enclose a branchial cavity within 
which lies the large, pedunculate, cup-like epipodite of the first thoracic limb 
(maxilliped) . Each lateral flap of carapace deeply separated anteriorly from the 
dorsal part ; cervical furrow visible in the better-preserved specimens. Thoracic 
somites 2-8 free (although the second is almost entirely overlapped by the carapace) , 
deepening progressively posteriorly. Abdomen long, exceeding half the total length 
of body ; telson distinct from the last somite. Ocular lobe oval, plate-like, movably 
articulated to side of rostrum, without visual elements or pigment. Antennulae 
almost contiguous basally, long, with two unequal or subequal flagella and a 3- 
segmented protopodite without statocyst but modified in the adult male. Antenna 
longer and more robust than antennula ; protopodite 2-segmented (peduncle 
therefore 4-segmented), exopodite small, scale-like, multiarticulate flagellum nearly 
as long as body. Mandible with lacinia mobilis, a series of 16-20 spines and a 
slender, 1 -segmented palp. Maxillula with a slender palp near distal end of outer 
margin of endite 3 (broad outer lobe). Maxilla well developed ; endite 3 deeply 
bifurcate, each lobe with long curved apical setae ; a few stout penicillate processes 
on endite 2 (inner lobe). Maxilliped " Isopodan " in form, with a few retinacula 
on inner margin of inner plate (endite of basipodite) a 5-segmented palp, no exopodite, 
a large respiratory cup-like epipodite. Lower lip without movably articulated 
apical lappets. Peraeopods simple, ambulatory, none markedly modified ; 
epipodites absent ; exopodites present on 1-6 (a rudiment on 7 is exceptional). 
Three anterior pairs of exopodites 2 (3) -segmented, setose, natatory ; three posterior 
pairs 1 (2)-segmented, non-setose, respiratory (gills). Pleopods alike in both sexes ; 
first four pairs well-developed, biramous, natatory, fifth pair vestigial. Uropods 
broad, biramous ; exopodite 2-, endopodite i-segmented. A simple penial process 
on coxopodite of peraeopod 7 in male ; incipient oostegites on peraeopods 2-5 in 
female (not mature) . In a more mature female sent after this paper was completed 
oostegites are present on peraeopods 1-5 inclusive (see p. 44) . 

Nothing is known of the internal anatomy or of the development. 

Gender of genus : masculine. Genotype : Spelaeogriphus lepidops n. sp. 

Holotype, allotype and the paratypes described below will be incorporated in the 
British Museum Collection. 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 33 

Spelaeogriphus lepidops n. sp. 

Description. The slender elongate body recalls that of many small cavernicolous 
Malacostraca ; it is subcylindrical being slightly depressed especially in the posterior 
third. In dorsal aspect the sides are almost parallel throughout, except posteriorly 
since the free telson is narrower than the abdominal and thoracic somites. In 
lateral aspect the body is as represented in Text-fig. i but the delicate side plates 
(pleura or epimera) of abdominal somites 1-4 are not at first glance apparent and 
therefore these somites appear to be less deep. 

The small, distinct carapace is smooth except for the cervical furrow which is 
distinct in the holotype, but only faintly indicated in the more transparent specimens. 
It is produced anteriorly, between the pair of oval ocular lobes or scales, to form a 
somewhat depressed, broadly triangular rostrum (Text-fig. 2). In lateral aspect, 
the carapace is as deep as long and, in all the preserved specimens (which may be 
somewhat contracted), it overlaps the first free thoracic somite (number 2) leaving 
only a small portion exposed dorsally, and part of the third somite laterally (Text-fig. 
1). Each lateral part of the anterior margin of the carapace is continued backwards, 
on a level with the outer rim of the ocular scale, for a considerable distance before 
fusing with the dorsal portion at the cervical furrow. Thus these antero-lateral 
flaps are doubtless capable of considerable lateral movement. Near the postero- 
lateral margin there is a conspicuous oval patch, above thoracic somite 2, represented 
by stippling in Text-fig. 1. This area, whose significance is unknown, is part of the 
carapace wall and can also be seen from the inside, as shown in Text-fig. za, where 
the ventral rim of the carapace is indicated, slightly posterior to the respiratory 
cup-like epipodite of the maxilliped. This large " gill " is visible through the thin 
wall of the carapace, but the oval patch behind it is not equally well marked in all 
the specimens ; for example, it is rather faint in the immature male paratype. 

Thorax. The first somite is completely fused with the head region ; the second 
somite is free from, but almost entirely overlapped by, the carapace. Somites 3-8 
become progressively deeper as represented in Text-fig. 1 although their dorsal 
margins are approximately equal in length. 

The abdomen exceeds half the total length of the body. Somites 1, 2 and 5 are 
subequal in length and shorter than the remaining three. Somites 1-4 decrease 
gradually in depth but, as already mentioned, their pleura are delicate and not 
easy to discern. The small epimeral portion of somite 5, however, is distinct 
(Text-fig. 1). 

The telson is free from, and narrower than, the sixth abdominal somite (Text-fig. 
16). The median length is nearly equal to the basal width and the rounded apex 
bears a number of spines of varying length. 

The antennulae, which are not widely separated from each other, are shorter and 
much less robust than the antennae. The proximal part of the right antennula 
of a female paratype is represented, in dorsal aspect, in Text-fig. 2, that of the 
holotype in later o- ventral aspect, in Text-fig. 3. The first segment of the protopodite, 
which is equal to the sum of the second and third segments, has no statocyst. In 
the adult male the second segment of the peduncle is modified, the distal half of the 
inner margin being expanded to form a lobe which is richly beset with rows of 



34 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 



05mm. 








0-5mm. 



Figs. 1-5. — Spelaeogriphus lepidops 11. g. and sp. Fig. 1. $ paratype, in lateral aspect, 
peraeopods omitted except for the exopodites or " gills " on 4-6 respectively. la. 
Ventral margin of carapace and cup-like epipodite at base of the maxilliped. Fig. 2. 
Lower figure — rostrum, right ocular lobe and proximal segments of antennula and 
antenna of a $ paratype, in dorsal aspect ; upper figure — rostrum and right ocular lobe 
of the smallest paratype, an immature $, in dorsal aspect. Fig. 3. Proximal segments 
of antennula and antenna of the holotype, in ventro-lateral aspect (the segments of the 
protopodite are broader than shown in figure) and, at higher magnification, a few distal 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 35 

conical papillae. Ventrally, this lobe is bounded by a series of long setae as shown 
in Text-figs. 3 and 4. There is also a small patch of the special, sharply pointed, 
conical papillae on the inner distal margin of the basal segment of the male peduncle. 
The longer flagellum comprises 40 or more segments, the smaller up to 36 segments ; 
in some specimens the two flagella are nearly equal in length. The distal three- 
fourths of the shorter (outer) flagellum has a series of short aesthetascs as shown in 
the enlarged portion (Text-fig. 3). 

The antennal peduncle of a female is represented in dorsal aspect in Text-fig. 2, 
that of the male holotype in latero-ventral aspect and at a lower magnification, in 
Text-fig. 3. There is no obvious sexual dimorphism of the peduncle or protopodite. 
The small, scale-like exopodite bears some 10 marginal spines. The flagellum when 
complete is almost as long as the body and comprises about 70 segments. 

The ocular lobe is a thin oval scale, movably articulated by means of a short 
peduncle to the side of the rostrum (Text -fig. 2) ; on the posterior outer edge is a 
tiny papilla. If viewed obliquely, the ocular scale seems to be decidedly club- 
shaped. There is no trace of pigment or of visual elements. 

The mouthparts represented in Text-figs. 8-14 are, with the exception of the 
maxilliped (Text-fig. 13), all from the same female paratype. The upper lip, as 
indicated in Text-fig. 1, is deep ; the distal portion is represented in lateral aspect in 
Text-fig. 8. The distal portion of the left mandible is shown (slightly distorted by 
pressure of the coverslip of the micropreparation) in Text-fig. 9. The palp is reduced 
to one elongate segment, the molar process is well developed and the spine row 
comprises 16-20 graded spines. There is only very slight asymmetry of incisor 
process and lacinia mobilis respectively (Text-fig. 10). The robust outer lobe 
(endite 3) of the maxillula bears about 16 short apical spines and, on the distal outer 
margin, a slender plumose seta-like palp (Text-fig. 11) ; the rather slender inner 
lobe (endite 1) has three short plumose terminal papillae. The maxilla, 
together with various details at a larger magnification, is represented in Text-fig. 
12 ; endite 3 is deeply bifurcate, the long curved spines on the outer lobe consist of 
a shaft and a slightly curved distal portion whereas those of the inner lobe have the 
apex bifurcate. The short inner spines of each series are somewhat conical with 
striate or serrulate concave (? outer) margins. Endite 2 has a few stout penicillate 
setae, three short and one long, on the inner margin. The maxilliped represented 
in Text-fig. 13 is from a considerably smaller specimen and, as it was not completely 
removed, the proximal portion (indicated by a broken line in the figure) may not 
be quite exact although the relative size and position of the epipodite is correct, 
the concavity facing inwards and forwards. The two maxillipeds are held firmly 
together by means of a few small retinacula on the inner margin of the inner plate 
(endite of the basipodite) . The palp appears to be 5-segmented ; the wide second 
segment bears 6 stout plumose setae on the inner margin ; the third segment, which 

segments of the shorter antennular flagellum. Fig. 4. Second segment of antennular 
protopodite of <$ paratype, in ventral aspect, to show specifically modified area beset 
with rows of conical papillae. Fig. 5. Vestigial pleopod 5 of q* paratype, highly magni- 
fied. Scale of Fig. 1 and ia = 1 mm. Scale of Figs. 2 (lower) and 3 respectively 
— 0-5 mm. 



36 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 




Figs. 6-14. Spelaeogriphus lepidops n.g. and sp. $ paratypes. Fig. 6. Distal segments 
of peraeopod 1 . Fig. 7. Exopodite of peraeopod 1 of a larger specimen. Fig. 8. Upper 
lip, in lateral aspect. Fig. 9. Distal part of left mandible, slightly distorted by the 
coverslip of microslide. Fig. 10. Portion of left and right mandibles, at higher magni- 
fication, to show slight asymmetry of incisor process and lacinia mobilis. Fig. 11. 
Maxillula, and apex of inner lobe more highly magnified. Fig. 12. Maxilla, with 
enlarged details of setae and spines. Fig. 13. Maxilliped of a smaller specimen, in 
ventral aspect (appendage broke just below the palp ; position of epipodite relative 
to the maxilliped is correct. Fig. 14. Lower lip. Figs. 8-12 and 14 from the same 
specimen. All except Fig. 10 drawn to scale = 0-5 mm. 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 37 

is contracted distally, also has marginal plumose setae ; the fourth segment is 
somewhat expanded distalfy and the fifth is inserted at an angle to the apex. The 
lower lip, represented in Text-fig. 14, shows no trace of the movably articulated 
lappet on each lobe which is characteristic of the family Apseudidae. 

Peraeopods. The peraeopods, which are simple and ambulatory (pediform), are 
arranged in two series, an anterior one of three, and a posterior one of four, pairs as 
shown in Dr. Barnard's sketch (Text -fig. 26). This division is emphasized by the 
fact that the exopodites on each of the three anterior pairs are natatory whereas 
those on peraeopods 4-6 respectively are reduced and modified to serve as gills 
(Text-figs. 1 and 18-20). In the allotype there is also a minute gill or exopodite on 
the right peraeopod 7, but this is exceptional. Each natatory exopodite consists of 
an elongate proximal, non-setose or sparsely setose, segment and a distal segment 
bearing marginal setae, the apical ones being especially long and often curled ; in 
addition a very short basal segment may be more or less clearly demarcated (Text-figs. 
7, 18, 21, 24 and 25). The respiratory exopodites, the posterior of which is quite 
small, lack the setose terminal segment ; the proximal segment is swollen and 
specially modified and again there is a more or less distinct small basal segment 
(Text-figs. 1, 19, 20 and 22). 

In the adult male peraeopods 1-3 are considerably more robust than in the female 
or young male, but otherwise there is no sexual dimorphism. The first peraeopod 
is the only one which may be regarded as slightly modified ; it is rather shorter than 
either peraeopod 2 or 3, with a smaller exopodite, a broader carpus, and more 
numerous spinules on the ventral margin of carpus and propodus respectively (Text- 
figs. 21 and 24). Text-figs. 24 and 25 represent peraeopods 1 and 3 respectively, 
drawn to the same scale, of the male paratype shown in Text-fig. 1 ; peraeopod 2 
is very similar to, though a trifle shorter than, peraeopod 3. In the holotype, which 
is rather larger, the spine-setae on the carpus are more numerous, especially in one 
of the two rows (3 and 5-6, as against 2 and 2 in the paratype represented in Text- 
fig. 25). When the distal segments are flexed these two rows lie one on either side 
of the propodus. Peraeopods 4-7 are relatively more slender and increase gradually 
in length owing chiefly to the progressive elongation of the carpus and especially 
of the propodus (cf. Text-figs. 22 and 23, peraeopods 4 and 7 respectively, of a male 
in the South African Museum). Peraeopod 7 as a rule lacks the exopodite ; a 
simple penial process is present in all three males on the coxopodite. This process is 
short in the smaller male paratype with the as yet unmodified antennular protopodite 
(and therefore rather immature) ; in the adult male it is long so that the two processes 
almost meet in the median line (Text-fig. 23) . 

The peraeopod represented in Text-fig. 6 was already detached from one of the 
paratypes but it probably came from one of the females ; in the allotype the first 
peraeopod has some spinules on the ventral margin of carpus and propodus and the 
exopodite is shorter, especially as regards the terminal segment, than that on either 
peraeopod 2 or 3 (cf. Text-figs. 7 and 18). Peraeopod 4 is more slender than peraeo- 
pod 3, with a longer more slender propodus ; the exopodite shows a hint of the short 
basal segment which, however, is distinct in that of peraeopod 5. What I consider 
to be incipient oostegites are present at the bases of peraeopods 2-5 but they are 



38 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 




0*5mk. 



Figs. 15-20. Spelaeogriphus lepidops n.g. and sp. Fig. 15. Uropod of allotype. 
Fig. 16. Telson and posterior margin of abdominal somite 6, in dorsal aspect. Fig. 
17. Fourth pleopod, with grappling hook more highly magnified. Fig. 18. Third right 
peraeopod of 2 with well developed natatory exopodite and incipient oostegite 0. 
Fig. 19. Fourth peraeopod with respiratory exopodite or " gill " and oostegite. Fig. 
20. Detached exopodite of peraeopod 5. Figs. 16-20 from same °- paratype. Figs. 
15 and 17 at smaller scale = 0-5 mm. ; rest at larger scale = 0-5 mm. 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 39 

not easy to make out (Text-figs. 18 and 19, 0). As in the male, peraeopods 5-7 
lengthen progressively and the respiratory exopodite on peraeopod 6 is smaller 
than that on peraeopod 5. Perhaps none of the female specimens is fully mature 
since the oostegites are so small and inconspicuous. 

Pleopods. These are alike in both sexes. The first four pairs are well developed 
and natatory. Each comprises a broad protopodite with two retinacula at the 
distal end of the inner, and a series of spines on the outer, margin and a paddle-shaped, 
heavily setose exopodite and endopodite (Text-fig. 17). The fifth pair is very much 
reduced and concealed by the small epimera of the fifth abdominal somite. In 
Text-fig. 5 the fifth pleopod has been displaced so that the drawing could be made, 
it is reduced to a slender club-shaped segment with 4 terminal setae. 

The uropods appear to be very fragile since in none of the specimens are they now 
complete ; either the exopodite, or both exopodite and endopodite are as a rule 
missing. In the vial containing the holotype and allotype two detached but complete 
uropods were found. These must belong to the allotype since in the holotype the 
peduncles are still attached to the specimen. One of these uropods is represented 
in Text-fig. 15 ; the short stout protopodite is furnished with an outer and an inner 
cluster of apical spines ; the exopodite is 2-segmented, the proximal segment is 
armed with spines along the inner margin and apically — the latter spines being 
especially long, the distal segment is very similar in shape and size to the exopodite 
and both are heavily setose. In the case of the male paratype represented in Text- 
fig. 1 the exopodite and endopodite had apparently undergone regeneration and are 
smaller than is normal for the species, so in the illustration I have indicated the 
normal length of the endopodite by a broken line and have omitted the exopodite 
(which in any case is most frequently missing). 

Remarks. The material available is insufficient for a study of the differences 
due to age and sex. Apart from that exhibited by the protopodite of the antennula 
(Text-figs. 2 and 3-4), sexual dimorphism appears to be slight. The segmentation 
of the antennular flagella is indistinct proximally, where the segments appear to be 
crowded, becoming more distinct distally. Dr. Barnard gives the number of seg- 
ments of the flagella as about 36 and 48 respectively but this is presumably the 
maximum number found in the largest specimen (I = 7*5 mm.). In the holotype 
the two flagella are not very unequal but the number of segments seems to be about 
32-33 and 40-42 respectively. In the allotype the flagella are markedly unequal 
yet each appears to have less than 30 segments (about 24 and 28 respectively). 
These differences may be due to sex. In the other specimens the flagella are not 
always complete. 

In the smallest male there is as yet no trace of the sexual modification of the 
first two segments of the antennular protopodite. Indeed, the antennula is relatively 
much shorter than in the adults, especially as regards the basal segment of the 
protopodite ; the smaller flagellum comprises only 15-16 segments. The larger 
flagellum is missing, as also are the antennae beyond the short protopodites (the 
two basal segments). The rostrum is also appreciably shorter than in the adults, 
and the ocular lobes are much more circular in outline (upper figure in Text-fig. 2). 



40 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 




0-5MM 



Figs. 21-26. Spelaeogriphus lepidops n.g. sp. Figs. 21, 22 and 23. Peraeopods i, 4 
and 7 respectively of a <J in South African Museum — sketches by Dr. K. H. Barnard ; 
magnification not stated. Specimen probably larger than the holotype. Figs. 24 and 
25. Left peraeopods 1 and 3 respectively, of <J paratype represented in Fig. 1. Scale 
= 0-5 mm. Fig. 26. Sketch of whole animal, in lateral aspect, by Dr. K. H. Barnard, 
showing arrangement of peraeopods in two groups, pleopods and uropod. 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 41 

The first peraeopod is very similar to that of the female (Text-fig. 6) ; the others are 
all incomplete although in most instances the proximal segments with the well 
developed exopodites remain. 

In a female paratype of approximately the same size as the male represented in 
Text-fig. i (/ = 6-5 mm.) the antennular protopodite is more slender and, especially 
as regards the first segment, shorter than in the male and there are fewer segments 
in the shorter flagellum ; the longer one is incomplete. The protopodite of the 
antenna is also relatively shorter and more slender (the male antennular and antennal 
protopodites are considerably broader than I have indicated in Text-fig. 3 where the 
appendages are lying obliquely, but I did not wish to risk damage to the holotype) . 
The first three peraeopods of the male become more robust with increase in size ; in 
the holotype they are somewhat more robust than in the paratype from which 
Text-fig. 24 was obtained and Dr. Barnard's sketch is probably from a male exceeding 
7 mm. in length. (Text-fig. 21). 

AFFINITIES OF THE GENUS S P E L A E OG R I P H U S 

In recent years many new cavernicolous and interstitial Crustacea Malacostraca 
have been discovered ; these are referable for the most part to the Orders Thermos- 
baenacea, Bathynellacea (Syncarida), Isopoda and Amphipoda (Peracarida). 

Spelaeogriphus, with its slender, elongate body, bears a striking resemblance 
to one of these cavernicolous forms namely, Monodella argentarii Stella (1951a, 
p. 2, fig. 1). But, the general similarity of telson, uropods, mandible (the palp 
excepted) and the exopodites 1 of the peraeopods notwithstanding, Spelaeogriphus is 
most certainly not referable to the Thermosbaenacea. This Order is unique amongst 
Malacostraca in the possession of a dorsal marsupium or brood-pouch in the female, 
" a chamber formed by the posterior portion of the carapace, which covers the first 
three somites of the body " (Stella, 19516, fig. 3 of plate ; 1953, pi. 1, fig. 2). Barker 
exhibited some ovigerous or larvigerous females of Thermosbaena mirabilis Monod, 
with a similar dorsal marsupium, at the XIV International Congress of Zoology 
held in Copenhagen in 1953, but his description and figures have not so far been 
published (Barker, 1953). Fertilization in the Thermosbaenacea must, therefore, be 
internal. In Monodella, according to Stella (1955, p. 464), from each ovary a short 
duct, the vagina, leads to the base of the sixth peraeopod and another one, the 
oviduct, goes dorsally to the brood-pouch. The position of the vaginal openings 
on the seventh thoracic somite (bases of peraeopods 6) is unusual ; in Malacostraca 
the female genital openings are, as a general rule, on the sixth somite. Another 
unusual feature in Monodella is the presence in the male of an additional coupling 
organ on the maxilliped (Stella, 1955, p. 464; Karaman, 1953, figs. 7 and 10). 
These characters, together with a study of the embryology of Monodella, led Taramelli 
(1954) to exclude the Order Thermosbaenacea from the Division Peracarida and 
with this Siewing agrees (1956, p. 168, Diagram 3). The Thermosbaenacea have 
certain characters of the Syncarida, others of the Peracarida, and still others which 
are unique. 

When Barnard first examined the specimens of Spelaeogriphus he thought that 

1 None of these exopodites are respiratory in Monodella. 



42 A NEW CAVERN I COLO US CRUSTACEAN FROM SOUTH AFRICA 

they belonged to the Division Syncarida. But, he writes, " further consideration 
shows the impossibility of including this Crustacean in that Division. Barring a 
superficial resemblance in having exopods on six of the peraeopods, it has none of 
the special features found in the Syncarida. On the contrary, its affinities seem to 
be with the Tanaidacea, especially Apseudes. The mouthparts are Isopodan in 
character, and the cup-like epipod on the maxilliped is clearly analogous to that 
found in Apseudes ' ' (Barnard, in letter received 14 . iv . 56) . After I had described 
Spelaeogriphus and had considered its possible relationships, I sent some notes and 
sketches to Dr. K. Lang, Director of the Stockholm Museum, since he has for some 
years past been engaged on a revision of the Order Tanaidacea. He replied in the 
following few words : " The animal you picture does not belong to the Tanaidacea 
but to the Anaspidacea " (letter dated 20.vii.56). Thus two eminent authorities 
on the lower Eumalacostraca disagree as to the systematic position of Spelaeogriphus. 

In my opinion Spelaeogriphus does not agree with either the Anaspidacea or the 
Tanaidacea as at present defined. In fact, like the Thermosbaenacea, it does not 
quite fit into either the Syncarida or the Peracarida. In Kiikenthal & Krumbach's 
Handbuch der Zoologie, Zimmer (1927, p. 566) defines the Divisions of the Eumalaco- 
straca and, as regards the external characters, Spelaeogriphus differs from the 
Syncarida and agrees with the Peracarida in having : (i) a carapace which encloses 
gill chambers but leaves most of the thoracic somites free ; (ii) a lacinia mobilis on 
the mandible ; (iii) two, not three, segments distal to the " knee " of the peraeopods ; 
(iv) oostegites in the female (but see later, p. 44). The antennal protopodite 
consists of two, not three, segments so that the peduncle comprises four segments ; 
this holds for some Syncarida and also (though not mentioned by Zimmer in his 
diagnosis on p. 566) for the Tanaidacea alone amongst the Peracarida (Caiman, 
1909, p. 191 ; Zimmer, 1927, p. 686). Thus there is something to be said in favour of 
Barnard's view that Spelaeogriphus is a primitive Apseudid. 

On the other hand, the genus differs in quite a number of respects from Zimmer' s 
(1927, p. 685) diagnosis of the Tanaidacea. There are seven, not six, free thoracic 
somites. The sides of the carapace are deep and separated anteriorly for a long 
distance from the dorsal or median part. The telson is distinct from, not fused with, 
the last abdominal somite. The abdomen itself is far longer than that of the Tanaids; 
but this difference may not be significant since in the Thermosbaenacea Monodella 
has a long, Thermosbaena a short, abdomen. The exopodites are more numerous 
and well developed, three pairs being natatory, three pairs respiratory ; in the 
Tanaidacea vestigial exopodites are sometimes present on the first two pairs of 
peraeopods only. While the form of the ocular lobe and of the epipodite on the 
maxilliped strongly recall the Apseudidae, Spelaeogriphus differs from that family 
in other respects, namely : The first peraeopods are not chelate or subchelate, nor 
is the second pair modified and fossorial. The antennulae are set closer together 
and are decidely smaller than the antennae. The mandibular palp is reduced to one 
segment, not " triarticulate " ; but this is probably of slight importance since the 
palp is absent in the family Tanaidae. The palp of the maxillula is not large and 
reflexed into the gill chamber, but small and placed near the distal end of endite 
3 (in Anuspides the palp is even smaller, though more proximally placed, see 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 43 

Chappuis, 1927, p. 596, fig. 584). There are no apical lappets on the lower lip. 
The uropods are broad, natatory whereas in Apseudids exopodite and endopodite 
are, as a rule, slender and multiarticulate. The Tanaidacea are entirely marine. 
I do not think that Spelaeogriphus is referable to the Order Tanaidacea, nor can it 
be placed in any of the other Peracaridan Orders — Cumacea, Mysidacea, Isopoda 
or Amphipoda. 

In addition to the characters already mentioned (p. 42), Spelaeogriphus differs 
from the Anaspidacea in other respects : Epipodites are absent from all the peraeo- 
pods, whereas in Anaspidacea there are one or two on each, with the exception of 
the last pair. The pleopods are alike in both sexes, and the endopodite is well 
developed in the anterior four pairs ; in the Syncarida the endopodite is rudimentary 
or absent, with the exception of the first two pairs in Anaspididae and Koonungidae, 
in which the endopodites are modified as copulatory organs in the male (Chappuis, 
1927, p. 594 ; Smith, 1909, figs 29 and 52 ; Nicholls, 1931, pi. 32, figs. 12 and 13). 
A thelycum or spermatheca appears to be absent in Spelaeogriphus but is present in 
Anaspidacea (Smith, 1909, fig. 27 ; Nicholls, 1931, p. 476, figs A and B). There is no 
statocyst in Spelaeogriphus such as occurs in e.g. Koonunga (Smith, 1909, p. 502, 
fig. 5). Certain characters of Spelaeogriphus, on the other hand, do recall those of 
some Anaspidacea. For example, both Spelaeogriphus -and Koonunga exhibit sexual 
dimorphism of the antennulae although the modified area in the male differs 
in position and in form in the two genera (c.f. Zimmer, 1927, p. 595, fig. 580 with 
Text-fig. 3 of the present paper). The three pairs of respiratory exopodites on 
peraeopods 4 to 6 in Spelaeogriphus are unusual — they resemble epipodites but from 
their position on the limbs both Barnard and I think they must be exopodites. 
The only other Eumalacostraca with exopodites of this type are Syncarida ; in 
the Anaspididae peraeopod 6 (thoracic limb 7) bears, in addition to the two epipodites, 
a reduced respiratory exopodite, whereas those on the anterior peraeopods are 
long and multiarticulate (Smith, 1909, p. 516, fig. 24 and p. 513, fig, 21). The free 
second thoracic somite, free telson and broad uropod recall the Anaspidacea and in 
Koonunga there is a distinct V-shaped notch above the attachment of the antenna 
in the frontal margin of the cephalon (Sayce, 1908, pi. 1, figs. 1 and 3) ; in Spelaeo- 
griphus there is a long slit in this position (Text-figs. 1, 2). It is possible to imagine 
a Syncarid with a carapace since, in the Division Peracarida, the carapace is present 
or absent and, when present, varies greatly in relative size. According to Barnard 
the mouthparts are Isopodan in character but the maxillula is not unlike that of 
Anaspidacea, especially the position and direction of the palp (Sayce, 1908, pi. 1, 
fig. 12 ; Smith, 1909, p. 508, figs. 13, 14). In the Anaspididae the mandible shows a 
hint of bifurcating although there is no lacinia mobilis, and the proximal epipodite 
on the maxilliped is large although not cup-like (Smith, 1909, figs. 9, 10 and 19). If 
Spelaeogriphus is a Syncarid it certainly is not referable to either the Anaspididae 
or the Koonungidae. Nor can it be placed with the minute rather degenerate 
members of the Bathynellidae although, if Ueno's observations are correct, some 
species of this family would seem to possess oostegites. Dr. Chappuis, whom I 
consulted on this point, writes "No! there is no brood pouch in Bathynella or 
Parabathynella ; the eggs are laid one after the other just where the animal happens 



44 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 

to be" (letter dated 22. v. 26). Yet Miuri and Morimoto (1953, p. 239) say of 
Bathynella morimotoi Ueno " Adult females carrying eggs and newly-hatched larvae 
are obtainable at all seasons of the year ". In the following year Ueno (1954, p. 
525, fig. 36) figures a long elliptical lamella on the coxopodite of the second peraeopod 
of Bathynella inlandica n. sp. and says that these structures, which are also present 
on the first pair of peraeopods, are presumably oostegites (marsupium). Here 
then is a Peracaridan character in certain species of the Bathynellacea. 

Like the Thermosbaenacea, Spelaeogriphus possesses certain characters of the 
Syncarida, others of the Peracarida. For the present it seems advisable to refer it 
to a new family, Spelaeogriphidae, with the characters of the genus, and to leave the 
systematic position of the family as uncertain. Perhaps when the internal anatomy 
and the embryology of Spelaeogriphus are known the systematic position of the 
family will be elucidated. As new forms of primitive Eumalacostraca come to light 
it may be necessary to revise the classification and even to redefine the major 
Divisions. 



ADDITIONAL NOTE 

After the manuscript was finished I received from Dr. Barnard two further 
specimens accompanied by the following note : " New species of shrimp ; pair 
found copulating. Bats Cave, stream at bottom. Collected by S.A.S.A. 29.7.56." 
The male and female were thought to be copulating when caught, and each should 
therefore be sexually mature. Unfortunately, uropods, antennulae and antennae 
are incomplete in both and in the male the posterior two or three peraeopods are 
broken and most of the gill-like epipodites are missing. 

In the male, which measures 6 ■ 5 mm. in length, the modified lobe on segment 2 
of the antennular protopodite is more pronounced distally than that represented in 
Text-fig. 4 and the conical papillae extend almost to the proximal articulation of 
the segment ; the patch on the inner distal margin of segment 1 is conspicuous. 
There seem to be a few papillae at the inner distal angle of segment 2, and a row of 
5 blunt cones on the inner margin of segment 3, of the antennal peduncle. The 
body of the female is slightly bent, but it appears to be rather shorter and is more 
slender than that of the male. The oostegites are quite unmistakable in this specimen 
although they are narrower than one might expect in a breeding female. In addition 
to the four pairs which I detected in the type specimens, a small pair is present on 
peraeopods 1. The first four pairs meet or even overlap medially ; each member 
of the fifth pair is only about as long as wide and does not quite reach the median line. 

In both specimens the peraeopods are rather bunched together and each is flexed 
towards its partner. Barnard sketches the peraeopods as arranged in two series, 
1-3 directed forwards and 4-7 directed backwards (Text-fig. 26) and in life this may 
be the case. 

There can now be no doubt as to the presence of a ventral thoracic marsupium 
such as is characteristic of the Division Peracarida. Spelaeogriphus, therefore, 
seems referable to that Division and, as far as the external characters are concerned, 
it agrees with the definition of the Peracarida given by Caiman (1909, p. 149) and 



A NEW CAVERN1COLOUS CRUSTACEAN FROM SOUTH AFRICA 45 

also that given by Zimmer (1927, p. 566) if very slightly modified to read "... Anten- 
nenstamm 2- oder 3-gliederig." (As already mentioned, Zimmer failed to recall that 
the antennal protopodite of the Tanaidacea is only 2-segmented, although he does 
mention this in his treatment of the Order on p. 686). However, the family Spelae- 
ogriphidae cannot be placed in the Order Tanaidacea for reasons which I have 
already given (p. 42). Nor can it be placed in any of the other Peracaridan Orders 
although the elongated abdomen and free telson, the large number of exopodites, 
and the sexual modification of the antennula in the male are characters which it 
shares with the Mysidacea. The only alternative, therefore, is to establish a new 
Order Spelaeogriphacea, with the characters of the family, to receive it. 

It is to be hoped that ovigerous females and larval stages may soon be collected and 
also that specimens fixed in Bouin or another suitable fixative will be available for 
sectioning. 



Postscript 18.xii.56. 

After the manuscript had gone to press I received five additional specimens 
collected in Bats Cave, on g.ix.56 by the S.A.S.A. One of these is an ovigerous 
female with a relatively large brood-pouch containing about 10-12 large ova ; the 
outlines of the separate oostegites are not clearly distinguishable but it is a normal 
Peracaridan brood-pouch composed, as already stated, of five pairs of oostegites 
on peraeopods 1-5 (somites 2-6). 

I also sent tracings of the illustrations to Dr. Rolf Siewing of Kiel, who has done 
some excellent work on the comparative morphology of the Crustacean Malacostraca. 
He replied as follows : " Mit grosser Freude habe ich Ihre Zeichnungen von Spelae- 
ogriphus lepidops studiert. Der neue Fund hat mich sehr interessiert . . . Meine 
Meinung nun zu der Neuentdeckung ist, dass es sich nicht um einen Vertreter der 
Syncarida handelt. Es fehlen bei Spelaeogriphus Epipodite, die bei den Syncarida 
wenigstens an einigen Thorakalextremitaten ausgebildet sind. Ein freier Carapax 
ist bei den Syncarida ebenfalls niemals ausgebildet. Die Oostegite und die Lacinia 
mobilis sind aber ganz typische Charakteristica der Division Peracarida. Ich halte 
es nicht fur wahrscheinlich, dass sich diesse Organe unabgehangig in einer anderen 
Kategorie der Malacostraca noch einmal entwickelt haben. Auffallig sind aber 
manche Ubereinstimmungen mit den Thermosbaenacea : Bau der Extremitaten 
des Thorax, Carapax, und Lacininia mobilis der Mandibel. Moglicherweisse ist 
Spelaeogriphus mit ihnen naher verwandt und stent ein primitives Bindeglied dar. 
Sicher wird die Untersuchung der inneren Anatomie weitere Aufschlusse geben." 

I too had been much impressed by the resemblances between Spelaeogriphus 
and Monodella — apart from the position of the marsupium, which is ventral in 
Spelaeogriphus and all the Peracarida, dorsal in the Thermosbaenacea. Dr. Siewing's 
comments have been most helpful and give me more confidence in proposing the 
new Order Spelaeogriphacea. The position of this primitive Peracaridan Order in 
Siewing's Diagram 3 (1956, p. 168) would appear to be within the Division Peracarida, 
near the suggested position of the Thermosbaenacea, thus : — 



46 A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 



CUMACEA 



AMPHIPODA 



DIVISION 
PERACARIDA 



SPELAEOGRIPHACEA 




THERMOS&AENACEA 



Marsupium 
jfcr- dorsal. 



With lacinia mobilis; 
/ marsupium thoracic. 



Adapted from Slewing, 1956, p. 168. Upper left-hand portion of Diagram 3. 



The Thermosbaenacea and the Peracarida have a lacinia mobilis on the mandible 
and a thoracic brood-pouch or marsupium. The Spelaeogriphacea have three pairs 
of exopodites modified as gills ; this may be a secondary specialization, perhaps an 
adaption to the freshwater habitat, although no other cavernicolous Malacostracan 
possesses such gills. The relationship of the Spelaeogriphacea to the other Peracaridan 
Orders must, for the present, remain uncertain. 



REFERENCES 

Barker, D. 1956. The morphology, reproduction and behaviour of Thevmosbaena mirabilis 

Monod. Proc. XIV Intern. Zool. Congr. Copenhagen : 503-504. 
Calman, W. T. 1909. Crustacea in Lankester's A Treatise on Zoology VII. London. 346 

pp., 194 text-figs. 
Chappuis, P. A. 1927. Crustacea Malacostraca — Syncarida in : Kukenthal & Krumbach's 

Handbuch der Zoologie. Bd. 3, Hft. 1 : 593-606, with text-figs. 
Jaeger, E. G. 1955. A Source Book of Biological Names and Terms. 3rd Editn. Springfield, 

Illinois. 
Karaman, S. 1953. Ueber einen Vertreter der Ordnung Thermosbaenacea . . . Mono- 

delta halophila n. sp. Acta Adriatica. Split, 5 (3) : 1-22, 24 figs. 
Mat j a sic, J. 1956. Em neuer Hohlendecapode aus Jugoslawien. Zool. Anz. 157 (3-4) : 

65-68, 2 text-figs. 
Miuri, Y. & Morimoto, Y. 1953. Larval development of Bathynella morimotoi Ueno. Annot. 

Zool. Jap. 26 (4) : 238-245, 4 text-figs., 2 tables. 



A NEW CAVERNICOLOUS CRUSTACEAN FROM SOUTH AFRICA 47 

Nicholls, G. E. 1931. Micraspides calmani, a new Syncaridan from the West Coast of 

Tasmania. Journ. Linn. Soc. Zool. 37 (no. 254) : 473-488, figs, a, b and pis. 31 and 32. 
Sayce, O. A. 1908. On Koonunga cursor, a remarkable new type of Malacotracous Crustaceans. 

Trans. Linn. Soc. London, Zool. (2) xi, 1 : 1-15, 2 pis. 
Siewing, R. 1956. Untersuchungen zur Morphologie der Malacostraca (Crustacea). Zool. 

Jahrb. Anat. Ontog. 75 (1) : 39-176, 72 text-figs. 
Smith, G. 1909. On the Anaspidacea., living and fossil. Q. J . Microsc. Sci., London, 53 

(3) • 489-578, 62 text-figs., pis. 11 and 12. 
Stella, E. 1951a. Monodella argentarii n. sp. di Thermosbaenacea (Crustacea Peracarida) 
Arch. Zool. I ted. Torino, 36 : 1-15, 22 figs. (Reprint.) 

1951k- Notizie biologiche su Monodella argentarii Stella . . . Boll. Zool. Torino, 18 

(4-6) : 227-233, 4 figs. 

1953- Sur Monodella argentarii Stella, espece de Crustace Thermosbenace . . . Hydro- 

biologia, 5 (1-2) : 226-232, 1 pi. 

1955- Behaviour and development of Monodella argentarii Stella, a Thermosbenacean 

from an Italian cave. Proc. Intern. Assocn. Limnol. 12 : 464-466. 

Taramelli, E. 1954. La posizione sistematica dei Thermosbenacei quale risulta dallo studio 
anatomico di Monodella argentarii Stella. Monitore zool. ital. Firenze, 62 (1) : 9-24, 3 pis. 

Ueno, M. 1954. The Bathynellidae of Japan (Syncarida-Bathynellacea). Arch. f. Hydrobiol. 
49 (4) : 519-538, 9 text-figs., 2 tables. 

Zimmer, C. 1927. Parts on Crustacea Malacostraca in : Kiikenthal & Krumbach's Handbuch 
der Zoologie. Bd. 3, Hft. 1 : 553-566 ; 683-696, with text-figs. 




PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON. LIMITED, 
BARTHOLOMEW PRESS. DORKING 




THE PELECANIFORM CHARACTERS 

OF THE SKELETON OF THE 

SHOE-BILL STORK, 

BALAENICEPS REX 



PATRICIA A. COTTAM 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 3 

LONDON: 1957 



THE PELECANIFORM CHARACTERS OF THE 

SKELETON OF THE SHOE-BILL STORK, 

BALAENICEPS REX 



BY 

PATRICIA A. COTTAM 



Pp. 49-72 ; Plate 3 ; 4 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 3 

LONDON: 1957 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5, No. 3 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued July, 1957 Price Eight Shillings 



THE PELECANIFORM CHARACTERS OF THE 

SKELETON OF THE SHOE-BILL STORK, 

BALAENICEPS REX 

By PATRICIA A. COTTAM 

CONTENTS 

Page 

Introduction . . . . . . . . . . 51 

Historical Note . . . . . . . . . 51 

Methods ............ 54 

osteological characters ......... 55 

A. Skull ............ 55 

b. Sternum and pectoral girdle ........ 62 

c. Pelvic girdle .......... 64 

d. Hind limb ........... 64 

Some Non-skeletal Pelecaniform Characters ..... 66 

Discussion ........... 66 

Summary ............ 70 

References ........... 71 

Plates ............ 72 

INTRODUCTION 

In the course of rearranging the bird skeletons in the collections of the British Museum 
(Natural History) it seemed to me that the skeleton of Balaeniceps rex had more 
pelecaniform than ciconiiform characters. The position of Balaeniceps in orthodox 
classifications is, and nearly always has been, near the storks and herons, so that 
this anomalous impression of its affinities seemed to require detailed investigation. 
The results of this study are presented here. The skeletal characters of Balaeniceps 
rex have been reassessed in relation to stork-like and heron-like characters on the 
one hand, and pelican-like characters on the other. 

I am grateful to Dr. H. W. Parker and to Dr. F. C. Fraser for reading the 
manuscript and making helpful comments, and also to Mr. J. D. Macdonald for 
advice and encouragement at all stages of the investigation. 

HISTORICAL NOTE 

There are few important original contributions to the knowledge of Balaeniceps s 
affinities. Gould, who described the bird in 1850, called it the " Grallatorial type 
of the Pelecanidae ", although he also noted that its external features resembled 
" in general contour " those of Grus, Ardea and Cochlearius. Jardine (185 1) noted 
likenesses to herons in the plumage. He considered that differences from the " true " 
pelicans, in the nostrils, pouch, position of the laryngeal opening and the absence 
of webs on the feet, were sufficient to show that Balaeniceps was not closely related 
ZOOL. 5, 3. 3 



52 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

to them. Von Heuglin (1856 : 60) placed it between Anastomus and Dromas in 
his systematic list. Bonaparte (1855 : 143) put it in the same subfamily as Coch- 
learius, describing it as intermediate between the pelicans and the Boat-bill. Des 
Murs (1859 : 4-8°) considered that the egg was like that of Phoenicopterus. 

These were the conflicting opinions in i860 when Parker examined the skeleton of 
Balaeniceps. He was impressed by its similarities to Scopus and Cochlearius, 
especially the latter, and indicated many characters which it had in common with 
the " Ardeine " birds. Although the storks were included in his term " Ardeine ", 
he seemed to stress the heron-like characters of Balaeniceps because he considered 
it to be a large edition of Cochlearius. He noted some similarities to the 
Pelecaniformes but attributed them to convergence. Bartlett's discovery (1861) 
of powder-down on Balaeniceps seemed to add weight to Parker's conclusions. 
Reinhardt (i860), unaware of Parker's work, found more similarities with Scopus 
than Cochlearius in the external characters of Balaeniceps, and considered that 
Balaeniceps and Scopus were nearer the storks than the herons. In 1861, after 
reading an abstract of Parker's paper, he compared a skull of Balaeniceps with those 
of Scopus and Cochlearius, but still maintained that Balaeniceps was related to 
Scopus and the storks. The similarities between the skulls of Balaeniceps and 
Cochlearius he attributed to convergence. Parker admitted (1862) that he knew 
nothing of the anatomy of Scopus when he wrote his paper, but, having seen a live 
Balaeniceps, he remained convinced of its likeness to the herons. He regarded 
Ardea as the " central type " of the storks and herons, linked to Cochlearius and 
Scopus by Balaeniceps. 

These two opinions became established. Some authors agreed with Reinhardt's 
conclusions and placed Balaeniceps with Scopus and the storks, but most of them 
agreed with Parker and placed it with Cochlearius and the herons. Giebel (1873) 
showed that the pectination of the middle claw and the pterylosis of Balaeniceps 
are similar to those of Scopus and different from Cochlearius. Beddard (1888) 
compared the visceral anatomy with that of the storks, herons and Scopus and, 
because of the alimentary tract, powder-down patches and syrinx, concluded that 
Balaeniceps was allied to the herons. Like Beddard, Furbringer (1888) and Gadow 
(1893 : 137) agreed with Parker. So did Shufeldt (1901), who wrote a paper on 
the osteology of Scopus and Balaeniceps without having seen a skeleton of the latter. 

The next important contribution was made by Chalmers Mitchell (1913) who 
dissected a specimen and described many more anatomical details. It is interesting 
that he could find no outstanding characters which indicated affinities with the 
herons rather than the storks, or vice- versa. He noted that Scopus and Balaeniceps 
had many similarities, and that they had characters common to both herons and 
storks. When he took each character at its face value he found Balaeniceps had 
more in common with the storks than the herons, so he decided to put it in the same 
suborder as Scopus, storks and ibises. He acknowledged that this was an arbitrary 
rather than a phylogenetic arrangement. He noted several similarities to the 
pelicans, but thought they occurred because the pelicans were related to the storks 
and herons. 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 53 

Bohm (1930) studied the structure of the skulls of juvenile and adult Balaeniceps. 
After a comprehensive investigation he concluded that Reinhardt's, Parker's, 
Giebel's and his own researches showed Balaeniceps to be a typical stork, linking 
the storks to the herons. He mentioned the " outstanding relationship " between 
Balaeniceps and Pelecanus, but did not seem to think it significant because he thought 
Pelecanus itself was so different from the other Pelecaniformes. The only other 
investigations of Balaeniceps 's anatomy were made by Technau (1937 : 567) during 
his studies of the nasal cavity of birds, and by Glenny (1955 : 560) in his work on 
the aortic arches of birds. The former drew no conclusions as to Balaeniceps s 
affinities, though Glenny thought it less like the Ciconiidae than is usually supposed. 

After Chalmers Mitchell's contribution most authors placed Balaeniceps by 
itself in a group of equal rank with the herons and storks (e.g. Streseman, 1927-34 : 
809 ; Wetmore, 1930 : 3). Mayr & Amadon (1951 : 6), however, followed Bohm's 
suggestion and placed it with the typical storks in the family Ciconiidae. Wetmore's 
classification (1951 revision) shows the generally accepted taxonomic position of 
Balaeniceps in relation to the orders Pelecaniformes and Ciconiiformes. 

Order Pelecaniformes. 

Suborder Phaethontes. 

Family Phaethontidae. 
Suborder Pelecani. 

Superfamily Pelecanoidea. 

Family Pelecanidae. 
Superfamily Suloidea. 
Family Sulidae. 

Phalacrocoracidae. 
Anhingidae. 
Suborder Fregatae. 

Family Fregatidae. 

Order Ciconiiformes. 
Suborder Ardeae. 

Family Ardeidae. 

Cochlearidae. 
Suborder Balaenicipites. 

Family Balaenicipitidae. 
Suborder Ciconiae. 

Superfamily Scopoidea. 

Family Scopidae. 
Superfamily Ciconioidea. 

Family Ciconiidae. 
Superfamily Threskiornithoidea. 
Family Threskiornithidae. 
Suborder Phoenicopteri. 

Family Phoenicopteridae. 
zool, 5, 3. 3 § 



54 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

METHODS 

As the purpose of this study is to examine the pelecaniform characters of 
Balaeniceps's skeleton in relation to its ciconiiform characters, the composition of 
the Pelecaniformes and Ciconiiformes will be discussed to decide what Balaeniceps 
ought to be compared with. 

The living members of the Pelecaniformes are apparently not very alike. Nearly 
every genus is placed in a separate family. From a comparison of their osteology 
it seems that the differences are mainly due to adaptive radiation, and that there 
is a well-defined basic similarity. For instance, superficially, pelicans and cormorants 
look less alike than storks and herons, but their skeletons have more characters in 
common. A possible exception is Phaethon, which is peculiar in many respects and 
may not be closely related to the rest of the Pelecaniformes. Wetmore (1951 : 5) 
thinks that " the Phaethontes possibly may have separated earlier than the 
Fregatae " from the pelecaniform stock. Therefore, as Phaethon is atypical the 
Phaethontes will not be referred to in this investigation. The Fregatae are also 
considered aberrant by some authors, but they have so many of the osteological 
characters typical of the Pelecani that they are probably fairly closely related 
to them. 

The Ciconiiformes is basically a less uniform group than the Pelecaniformes. 
Osteologically, it seems to be a collection of unrelated groups which, superficially, 
only have long beaks, long necks and long legs in common. 

The genera of the Ardeae are very alike, their outstanding variation being in 
size. Cochlearius is the most aberrant genus but, apart from its skull, it has all 
the characters of the typical herons. Even in its skull the heron-like characters are 
not completely obscured. For the present purpose, therefore, the Ardeidae and 
Cochlearidae will be considered together, as a monophyletic group representing 
the herons. 

The families in the Ciconiae are not so closely related. The Scopidae, with its 
single monotypic genus Scopus, is as enigmatic in its relationships as Balaeniceps. 
The skeleton of Scopus is like that of a small stork in some characters, but very 
unlike it in others. It has often been compared with Balaeniceps, and most authors 
consider the two related. However, there is no point in comparing one genus of 
doubtful affinities with another, so Scopus will not be referred to. The Ciconiidae is 
probably a monophyletic group ; its genera are fairly alike, although they vary 
more than those of the Ardeidae. This variation mainly seems to be due to different 
adaptations of the beak, correlated with differences in the size and shape of the head. 
The third family, Threskiornithidae, appears to have much in common with the 
Ciconiidae, but it also has certain resemblances to the Phoenicopteri. As the 
affinities of the Phoenicopteri themselves are controversial it is advisable not to 
discuss either group until their relationships have been more fully investigated. 

For the purposes of this investigation, therefore, Balaeniceps is compared with 
the suborders Pelecani and Fregatae, representing the Pelecaniformes ; the ciconi- 
iform suborder Ardeae, representing the typical herons ; and the family Ciconiidae, 
representing the typical storks. 



THE PELECANIFORM CHARACTERS OF THE SHOE-BIL STORK 55 

Three complete skeletons of Balaeniceps and three skulls were available. There 
was also adequate material of pelicans and their allies, frigate birds, herons and 
storks. The skeleton of Balaeniceps was systematically compared with those of 
Pelecanus, Ardea amd Ciconia, but other genera, especially in the Pelecaniformes, 
were consulted to determine the range of variation in each group. 

For convenience, in the following description Nannopterum, Halietor and Anhinga 
will not be mentioned unless they differ from Phalacrocorax. 



OSTEOLOGICAL CHARACTERS 

A. Skull, see Plate 3 

(1) Premaxilla 

Of the Pelecaniformes considered here, Pelecanus, Phalacrocorax and Fregata 
each have a well developed hook at the tip of the premaxilla. The newly hatched 
chick of Sula also has this hook, but it decreases with age, and in the adult the tip 
of the premaxilla is only slightly decurved. Anhinga has no hook in chick or adult, 
but this may be an adaptation to its habit of spearing fish. 

In the Ciconiidae there is no suggestion of a hook to the premaxilla in any of the 
genera. The nearest approach is the decurved bill of Ibis and Mycteria, but in 
these the distal fifth of the mandible is involved in the curvature. 

The Ardeidae have straight bills. Parker (1862 : 299) argues that in Cochlearius 
a hook " certainly does exist, although feebly " but, although the tip of the 
rhamphotheca is decurved, it is not hooked, and the premaxilla is quite straight 
ventrally. 

Balaeniceps has a prominent hook at the tip of the premaxilla, like the typical 
Pelecaniformes. 

(2) Nasal groove 

In the Pelecani and Fregatae there is a conspicuous groove running along each 
side of the culmen from the anterior edge of the nostril to the cutting edge of the 
premaxilla beside the terminal hook. This relationship of the nasal groves to the 
premaxillary hook is constant in Pelecanus, Sula, Phalacrocorax and Fregata. In 
Anhinga the groves are only faintly indicated. 

In the Ciconiidae the nasal groves are either absent, or represented by very shallow 
depressions which extend from the nostrils to, at most, half-way along the beak. 
Both conditions are often found in the same species. 

The Ardeidae have shallow depressions like those of the Ciconiidae instead of 
nasal grooves. In Cochlearius these depressions are expanded to form broad, 
shallow troughs, each with a ridge along the mesial border. 

Balaeniceps has conspicuous nasal groves which extend from the nostril to the 
cutting edge of the premaxilla beside the terminal hook, exactly as they do in the 
Pelecani and Fregatae. The grooves are not shallow, like those of the Ardeidae 
and Ciconiidae, or broad like those of Cochlearius, but deep like those of Pelecanus. 



56 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

(3) Nasal septum 

In the Pelecani and Fregatae there is an ossified nasal septum. The nasal septum 
ot the Ciconiidae and Ardeidae is not ossified, and it is perforated in the region of 
the external nares. Cochlearius has a complete, unossified nasal septum. In 
Balaeniceps the nasal septum is ossified, as it is in the Pelecani and Fregatae. 

(4) Nasal passage 

In Pelecanus the external nares are vertically above, or even slightly posterior 
to the internal nares, and the nasal cavity lies almost vertically between them. In 
the other Pelecani the external nares are only slightly anterior to the internal nares. 
In the Ciconiidae and Ardeae the external nares are an appreciable distance anterior 
to the internal nares, and the nasal cavity lies obliquely between them. In 
Balaeniceps the relative positions of the nares and nasal cavity are exactly the same 
as they are in Pelecanus. 

(5) Palate (See Fig. 1) 

In the Pelecani and Fregatae the palatines are always ankylosed along the mid- 
line posterior to the internal nares. There is usually a median ventral ridge, more 
or less well developed, along the suture, with a depression for the pterygoid muscle on 
either side of it. These depressions extend forwards past the posterior edge of the inter- 
nal narial opening only in Fregata. In the region of the internal nares the mesial 
edges of the palatines are parallel in the Pelecani, and nearly so in the Fregatae. In 
Pelecanus the ventral part of the nasal passage is divided along the mid-line by a 
membranous septum. There is no trace of an ossified pre vomer in association with 
this septum, and in Sula the septum itself is weakly developed. The septum is 
better developed in Phalacrocorax, and in at least one species, P. utile, there is a 
thorn-like cartilaginous prevomer associated with it (unless it is carefully dissected 
out the prevomer is easily lost in prepared skeletons of Phalactocotax) . In 
Fregata the prevomer is also thorn-like, though longer and definitely ossified. The 
maxillopalatines vary in size throughout the Pelecani and Fregatae. In Sula and 
Phalacrocorax they are small and do not project beyond the palatines mesially. 
They are slightly larger in Fregata, and can be seen, in ventral view, bordering the 
anterior half of the internal nasal opening. In Pelecanus they are very large and 
meet in the mid- ventral line between the anterior ends of the palatines. Also in 
Pelecanus, they nearly fill the inside of the skull in the nasal region, and their posterior 
edges slant backwards in a straight line from the internal nares to the cranio-facial 
hinge. Posteriorly, the maxillopalatines do not extend past the cranio-facial 
hinge-line in any of the Pelecani or Fregatae. 

The palatines of the Ciconiidae are not fused along the mid-line except at one point. 
Instead of the median ventral ridge found in the pelicans, there is a ventral crest along 
the mesial edge of each palatine where it borders the internal narial opening. The 
depressions for the pterygoid muscles extend further forward on either side of the 
narial opening than in Fregata. Immediately anterior to the internal nares the 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 57 





Fig. 1. Diagrams of ventral views of palatine regions of (a) Balaeniceps rex, (b) Pele- 
canus crispus, (c) Phalacrocorax urile, (d) Ciconia ciconia, (e) Ardea goliath, (f) Coch- 
learius cochlearius. 

1 = maxillopalatine, 2 = prevomer, or position of unossified septum, 
3 = palatine, 4 = depression for pterygoid muscle. 



58 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

palatines approach the mid-line, and in the larger genera, such as Leptoptilos and 
Jabiru, they may even touch. This divides the space between them into anterior 
and posterior parts, the nasal passage being confined to the posterior part. The 
prevomer arises at the posterior end of the internal narial opening, and the palatines 
are ankylosed at this point. The prevomer varies from a narrow strip of bone in 
Ibis, to a substantial triangular plate, drawn out into a thin filament anteriorly, 
in Leptoptilos and the larger genera. The maxillopalatines are well developed when 
compared with those of most Pelecani and Fregatae. They always meet in the mid- 
ventral line, where they occupy most of the space between the palatines anterior 
to the internal nares. Each maxillopalatine is extended posteriorly into a convex 
projection which usually reaches beyond the cranio-facial hinge ; in the Pelecani 
there is no such projection. 

The palatines of the Ardeidae are like those of the Ciconiidae except that they are 
separate along the mid-line, even where the prevomer arises. In Cochlearius, 
however, they are ankylosed at this point, as they are in the Ciconiidae. The 
depressions for the pterygoid muscles extend as far forwards on either side of the 
internal narial opening as they do in the Ciconiidae. In the Ardeae, unlike the two 
previous groups, the vomer is V-shaped in cross-section, though this is less obvious 
in Cochlearius. The maxillopalatines meet in the mid-ventral line anterior to the 
nasal opening, much as they do in the Ciconiidae. Their posterior edges are convex 
like those of the Ciconiidae, and extend well beyond the level of the cranio-facial 
hinge. The maxillopalatines are smaller in Cochlearius, but otherwise they are very 
like those of the Ardeidae. 

In Balaeniceps the palatines are ankylosed along the mid-line, posterior to the 
internal nares, with a broad ventral ridge along the suture. The depressions for 
the pterygoid muscles lie on either side of this ridge ; they extend forward to the 
level of the nasal aperture, but no further. The condition of the palatines is thus 
very like that of Pelecanus. The prevomer is weakly developed and its degree of 
ossification varies in the specimens examined. It is a thin, triangular plate, often 
perforated, lying in a vertical plane. In some specimens the apex does not reach 
the anterior end of the nasal opening. This weak development of the prevomer is 
reminiscent of the Pelecani, in which the prevomer is reduced and sometimes missing. 
The maxillopalatines are strikingly like those of Pelecanus. Their posterior faces 
are flat, even concave, and not convex like those of the Ciconiidae and Ardeae 
(Pycraft, 1898 : 83). 

(6) Lachrymal (See Fig. 2) 

The lachrymal in the Pelecani and Fregatae descends from the frontal to the 
quadrat ojugal bar, to which it is usually attached by a ligament. Viewed posteriorly, 
it is a column of bone with a lateral groove, of varying depth, to accommodate 
the lachrymal duct. In Phalacrocorax a narrow lateral process of the interorbital 
septum meets and fuses with the ventral end of the lachyrymal. This process 
is larger in Anhinga, lying beside the lachrymal throughout its length without touch- 
ing it. In lateral view, the lachrymal is more or less pillar-shaped in Pelecanus, 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 59 

Fregata and Phalacrocorax, but in Sula it is expanded anteriorly into the antorbital 
vacuity. There is a tendency, in the Pelecani and Fregatae, for this vacuity to be 
reduced. In Pelecanus it is comparatively large. In Fregata the maxilla grows 
back into it a little posteriorly. In Phalacrocorax there is a splint of bone resting 
on the quadrat ojugal bar. This bone fills most of the antorbital vacuity in Anhinga, 
in which the maxilla is produced posteriorly as well. The large lachrymal itself 
fills most of the antorbital vacuity in Sula, though the maxilla and the quadratojugal 
also expand into it. 

Although the lachrymal is well developed in the larger Ciconiidae, it never reaches 
the quadratojugal bar. In posterior view it is roughly triangular, with the apex 
of the triangle downwards. In some genera, including the four largest, the lachrymal 





/cm. 



I cm 



I cm. 



I cm. 



I cm. 



I cm. 



Fig. 2. Diagrammatic transverse sections of lachrymals of (a) Balaeniceps, (b) Sula, 
(c) Ciconia, (d) Leptoptilos, (e) Ardea, (f) Cochlearius. 
1 = lachrymal duct, 2 = lachrymal bone, 3 = quadratojugal bar. 



duct is wholly or partly enclosed in bone, giving a flat surface to the outer face of 
the lachrymal bone. Unlike the Pelecani or Fregatae, the ciconiid lachrymal has 
a mesial projection extending towards the interorbital septum and passing ventral 
to the duct of the nasal gland. The lachrymal is triangular in cross-section, and it 
never extends into the antorbital vacuity. This vacuity is large in the Ciconiidae, 
and there is no obvious tendency for the surrounding bones to expand into it. 

In the Ardeidae the lachrymal nearly reaches the quadratojugal bar. Its shape 
seems to be peculiar to the Ardeidae and is quite different from the Pelecani, Fregati 
and Ciconiidae. In Cochlearius the lachrymal is reduced, and in lateral view looks 
different from that of the Ardeidae ; but in cross-section it is almost identical. The 
antorbital vacuity is large in the Ardeidae and in Cochlearius. 

Balaeniceps has a large lachrymal. In posterior view it is like the lachrymal of 
the Pelecani and Fregatae, a column of bone which meets the quadratojugal bar 
ventrally. The lacrymal duct lies in a large foramen through the lachrymal bone, 
as it does in some Ciconiidae. Anteriorly, the lachrymal fuses with the maxilla, 
so that the antorbital vacuity is obliterated. There is a slight groove which may 



6o THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

represent the suture between the lachrymal and the maxilla. If it does, the lachrymal 
is pillar-shaped in lateral view as it is in the Pelecani and Fregatae. The complete 
occlusion of the antorbital vacuity, which occurs in Balaeniceps, is not found in 
any of the other groups considered here, but the Pelecani and Fregatae have a 
tendency towards reduction of the size of the antorbital vacuity. 



Icm. 





Icm. 





Fig. 3. Diagrams of articulating surfaces of quadrates and lower jaws of (a) Balaeniceps 
rex, (b) Sula bassanus, (c) Ciconia ciconia, (d) Cochlearius cochlearius. 

1 = left quadrate, 2 = left ramus of lower jaw, 3 = mesial articulating facets, 
4 = lateral articulating facets, 5 = lateral process. 



(7) Lower jaw articulation (See Fig. 3) 

Each of the three groups being compared with Balaeniceps has a different arrange- 
ment of the articulating surfaces of the quadrate and lower jaw. The arrangement is 
constant within each group, except that the one typical of the Ardeidae is found 
mainly in the larger species. 

In the Pelecani and Fregatae there are two articulating facets. On the quadrate, 
the mesial articulation has a broad ridge, which slides in a corresponding trough in 
the lower jaw. The long axis of the articulation lies at an angle of about 45 ° to the 
long axis of the skull, and is in line with the long axis of the pterygoid. This is 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 61 

especially noticeable in Pelecanus. In Sula, Fregata' and Phalacrocorax the lateral 
edge of this ridge on the quadrate is undercut, and the corresponding side of the 
groove in the lower jaw is overhanging. The result is a locking mechanism which, 
in the dried skull, is efficient enough to hold the lower jaw in place when the beak is 
closed. The lateral articulation is usually S-shaped, though in Sula it may be 
L-shaped. Its long axis lies approximately at right angles to that of the other 
articulation. In Pelecanus, possibly because of its wide gape, the lateral part of 
this articulation is modified. In the lower jaw, instead of a sigmoid articular surface 
there is a groove, running anteriorly, parallel to the mid-line. Along this groove 
slides part of the lateral articulating surface of the quadrate. This groove in the 
lower jaw is represented in Sula by a shallow transverse trough, the anterior side of 
which projects laterally and dorsally and lies anterior to the lateral process of the 
quadrate when the beak is closed. The lateral process on the lower jaw is reduced 
in Fregata and Phalacrocorax. 

The Ciconiidae also have two articulating facets. Unlike the Pelecani and Fregatae 
the long axis of the mesial facet is at right angles to the long axis of the skull, and at 
an angle of about 120 to the pterygoid. There is no locking mechanism. The 
lateral articulation is curved, so that while its lateral end is at right angles to the 
long axis of the mesial articulation, its mesial end is parallel to it. The lateral 
process on the lower jaw is well developed. The relationship between the two facets 
is quite different from that found in the pelicans or herons, and it is very alike in 
all the Ciconiidae examined, whatever the relative proportions of bill and skull. 

In most of the larger Ardeidae there are four articulating facets, as each of those 
occurring in the pelicans and storks is in two parts. The lateral part of the mesial 
facet and the mesial part of the lateral facet lie on a plane nearly parallel to the 
pterygoid. On the quadrate the mesial facet, although it is in two parts, is undercut 
laterally to give a locking mechanism, as it is in most of the Pelecani and Fregatae. 
The lateral process on the lower jaw is more prominent than in the other groups 
described. In Cochlearius the articulating facets are essentially the same as in the 
larger herons, but the mesial facet on the quadrate is undercut mesially as well as 
laterally, apparently increasing the efficiency of the locking mechanism. The 
lateral process on the lower jaw is even better developed than in the Ardeidae, and, 
with the lateral part of the lateral articulation, seems to function as an auxiliary 
locking device. 

Balaeniceps has two undivided articular facets, like the Pelecani, Fregate and 
Ciconiidae. On the quadrate, the mesial facet consists of a broad ridge, undercut 
laterally, which, in the lower jaw, slides in a trough with an overhanging lateral edge, 
much as it does in Sula, Phalacrocorax and Fregata. The mesial side of the trough 
also overhangs slightly, but not as much as in Cochlearius. The axis of the mesial 
articulation on the quadrate is in line with the pterygoid, as it is in the Pelecani 
and Fregatae, and is in contrast to the condition in the Ciconiidae. The lateral articula- 
tion is L-shaped, as it is in Sula ; its long axis is nearly at right angles to that of 
the mesial facet, like the Pelecani and Fregatae, and unlike the Ciconiidae. In 
the lower jaw, the lateral process is insignificant and the lateral articulation takes no 



62 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

part in the locking mechanism as it does in Cochlearius. Balaeniceps has none of 
the well defined ciconiid characters in its jaw articulation ; it resembles the Ardeae 
in some ways, but differs in others ; it is like the Pelecani and Fregatae in all the 
characters in which they differ from the Ciconiidae and Ardeae. 

b. Pectoral Girdle (See Fig. 4) 

(1) Furculum 

There is a tendency in the Pelecani and Fregatae for the hypocleideum of the 
furculum to be fused to the keel of the sternum. The joint is ligamentous in 
Phalacrocorax ; sometimes ankylozed in Sula ; usually ankylosed in adults of 
Pelecanus ; and so ossified in adults of Fregata that the suture is obliterated. 
Except in Fregata each arm of the furculum forms an arc, convex anteriorly, between 
each coracoid and the carina sterni. Characteristic of the typical pelecaniform 
pectoral girdle is the well developed acrocoracoid flange, which forms a flat transverse 
surface on the clavicle for articulating with the coracoid. In Fregata the clavicle 
is completely fused to the coracoid in this region, but in young specimens the presence 
of the acrocoracoid flange can be inferred from the sutures. Although an acro- 
coracoid flange is present in several other apparently unrelated groups, it is never 
as well developed as it is in the Pelecani, Fregatae, Balaeniceps and Scopus. 

In the Ciconiidae, although the hypocleideum of the furculum joins the carina 
sterni, it forms a bony fusion with it only in some specimens of one genus, Leptoptilos, 
and the suture is always obvious. Unlike the Pelecani, each clavicle forms a sigmoid 
curve in lateral view. The dorsal part of the curve is convex anteriorly, and the 
ventral part, where the clavicles unite in the mid-line is concave anteriorly. There 
is no indication of an acrocoracoid flange in any of the Ciconiidae. 

The furculum of the larger Ardeae is mainly like that of the Ciconiidae. The 
joint between the hypocleideum and the carina sterni is always ligamentous. The 
presence of a small interclavicle is characteristic of the Ardeae, and it is not found 
in the other groups considered here. 

The hypocleideum of Balaeniceps is fused to the carina sterni as it is in Fregata 
and most adult specimens of Pelecanus. The suture is obliterated by ossification 
in all the British Museum specimens. The clavicle of Balaeniceps forms a continuous 
arc from the coracoid to the carina sterni, as it does in the Pelecani. This character 
may not be significant, as the clavicle of Fregata is in a slightly sigmoid curve and 
that of Cochlearius is almost in a continuous curve. The acrocoracoid flange is 
well developed in Balaeniceps, a character typical of the Pelecaniformes. 

(2) Sternum 

In Balaeniceps the sternal keel extends along the whole length of the sternum to 
the posterior border as it does in the Ciconiidae, Ardeae and most other birds. In 
Pelecanus, Sula and Phalacrocorax it only reaches half-way back from the anterior 
end of the sternum. Parker (i860 : 329) considered this a significant difference 
between Balaeniceps and the Pelecani, but apparently was not aware of the condition 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 63 




64 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

in Fregata, in which the keel extends almost to the posterior end of the sternum. 
Thus within the Pelecaniformes both types of keel occur. 

c. Pelvic Girdle 

Parker (1861 : 336) considered the pelvis of Balaeniceps to be " typically ardeine " 
because it was narrow like that of the Ardeae. Chalmers Mitchell (1913 : 696) 
thought it more like the ciconiid pelvis because it had a notch in the posterior border, 
like the Ciconiidae, and lacked the ridge on the ilium which is present in the Ardeae. 
However, the shape and details of the pelvis in birds seem to depend mainly on 
the function and relative size of the legs and leg muscles, and the pelvis is probably 
a very adaptable part of the skeleton. In the Pelecaniformes, for example, the pelvis 
of Pelecanus, a bird with strong legs, seems to have very little in common with that 
of Phalacrocorax, in which the legs are weaker and used mainly for swimming, or of 
Fregata, in which the legs are very weak and only used for perching. In groups 
in which there is less adaptive radiation, such as the Ciconiidae, or Ardeae, the 
function of the legs is more uniform and the shape of the pelvis varies little within 
the group, the main differences being in size. In Balaeniceps the pelvis is roughly 
the same shape as it is in the Ardeae and some Ciconiidae, but it differs from both in 
details. It seems even less like that of any of the Pelecaniformes, but as there is 
already a good deal of variation of the pelvis in this group Balaeniceps would perhaps 
be less out of place with them than with the Ciconiidae or Ardeae. 

d. Hind Limb 
(1) Tibio-tarsus 

There are two forms of the distal condyles and the inter-condylar sulcus of the 
tibio-tarsus in the groups considered here. One is found in the Pelecani, Fregatae 
and Ardeae. In it the distal condyles are roughly semicircular in lateral view and 
the distal border of the outer condyle has no notch. The anterior aspect of the 
inter-condylar sulcus is fairly shallow, and the knob on the tarso-metatarsus which 
fits into it is not well developed. This type of articular surface is probably un- 
specialized, as it occurs throughout the Pelecani and Fregatae, in which there is 
considerable variation in the function of the legs, and in the Ardeae, in which the 
legs are long and unlike those of any of the pelican groups. 

A second form occurs in the Ciconiidae. In it the distal borders of the condyles 
are flattened, and the condyles themselves are elongated posteriorly, so that they are 
oval in lateral view. There is a notch in the distal border of the outer condyle. 
The anterior aspect of the inter-condylar sulcus is deep, and proximal to it there is a 
hemispherical depression with a prominent knob immediately beside it. The knob 
on the tarso-metatarsus is much larger than in the first type, and articulates with 
the hemispherical depression when the leg is bent. The second condition apparently 
only occurs in long-legged birds, such as the Threskiornithidae and Phoenicopteridae, 
and to a lesser extent in the Gruidae and long-legged Charadrii. 

The form in Balaeniceps is similar to that of the Pelecani, Fregatae and Ardeae. 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 65 

(2) Tarso-metatarsus 

In most groups of birds the hypotarsus is well ossified to form a varying number 
of " bridges " which enclose the flexor tendons in bony tubes. Of the Pelecani, 
Phalacrocorax has one tube and Sula and Pelecanus two. 

In contrast, the Ciconiidae have a so-called " simple " hypotarsus. It consists of 
two parallel bony ridges with a groove between. The flexor tendons lie in this 
groove and are supported by unossified ligaments instead of bony bridges. 

The Ardeae have a more ossified hypotarsus, rather like that of the Pelecani. 
In most genera there is only one tube, but the smaller genera sometimes have 
more. 

Balaeniceps has two complete bony tubes through the hypotarsus. Their 
arrangement is strikingly like that of Pelecanus, and quite unlike the Ciconiidae. 

(3) First metatarsal 

In the Ciconiidae and Ardeae the first toe points backwards. In the Pelecani 
it is joined to the second toe by a web and is restricted to a lateral position, although 
it is mobile enough to be able to be pointed forwards. Parker (1861 : 344) says 
that in Balaeniceps the first toe is " very mobile " and is turned " very far inwards " 
when walking. Photographs show that it is directed backwards when the bird is 
standing still. 

The position of the first toe influences the form of the first metatarsal. When the 
toe normally points backwards the metatarsal, if straight, would lie in the same plane 
as the flexor tendons of the other digits and interfere with their functioning. But 
the shape of the metatarsal is modified, usually giving it the appearance of bending 
round to one side of the tendons, and it often has a diagonal groove in which the 
tendons run freely. 

In Pelecanus, in which the first toe does not point backwards, the metatarsal is 
straight, with only a shallow depression, mid-way along its length, where it touches 
the flexor tendons. In the other Pelecani this depression varies in size and depth, 
but it is never so marked as it is in the Ciconiidae and Ardeae. In Sula and Phala- 
crocorax the metatarsal is slightly bent round the flexor tendons. 

In the Ciconiidae there is a broad, deep, diagonal trough for the flexor tendons, 
and the metatarsal appears twisted through an angle of about 90 . In the Ardeae 
the first metatarsal does not press against the flexor tendons as closely as it does in 
the Ciconiidae, because of the way in which it articulates with the first phalanx. 
As a result the diagonal groove in which the tendons lie is less marked than it is in 
the Ciconiidae and narrower than it is in the Pelecani. 

In Balaeniceps the metatarsal has a depression for the flexor tendons which is 
very little deeper than that of Pelecanus. It is shallower than that of the Ciconiidae 
and broader than that of the Ardeae. The metatarsal appears slightly twisted, 
though less so than it is in the Ciconiidae. The form of the first metatarsal and the 
function of the first toe of Balaeniceps therefore seem to be more like those of the 
Pelecani than the Ciconiidae or Ardeae. 



66 THE PELECANIFORM CHARACTERS OF THE SHOE-BIL STORK 

(4) Toe articulations 

The proximal articulating surfaces of the basal phalanges of the second, third 
and fourth digits are fairly alike in the Pelecani and Ciconiidae, being roughly square 
in shape. In the Ardeae each of these articulating surfaces has a characteristic, 
irregular shape. Balaeniceps is like the first two groups, with the articulations 
almost square in proximal view. 

SOME NON-SKELETAL PELECANIFORM CHARACTERS 
OF BALAENICEPS 

{a) von Heuglin (1873 : 1095) 

The egg is white with chalky lumps. Similar chalky lumps and nodules occur on 
eggs of Phalacrocorax and Sula. 

Birds join up in parties to herd shoals of fish into corners. This communal fishing 
is characteristic of some Phalacrocorax and Pelecanus species. 

(b) Chalmers Mitchell (1913) 

The rhamphotheca is compound, as it is in the Pelecani and Fregatae. 
A pyloric chamber is present in the stomach, as in Pelecanus. 
The dermo- temporalis, biventer maxillae, temporal and pterygoid muscles are 
similar in Pelecanus. 
There are no intrinsic muscles of the syrinx in Balaeniceps and Pelecanus. 
The hyoid muscles are " excessively like those of Pelecanus." 
The division of the pectoral muscle is similar in Pelecanus. 
The arrangement of the wing tendons is the same in Pelecanus. 

(c) Technau (1936 : 567) 

The secondary nostrils can be closed, as in Pelecanus. 

(d) Glenny (1955) 

The right carotid is absent in Balaeniceps and some Pelecaniformes. When one 
carotid is missing in the Ciconiiformes it is the left one. 

DISCUSSION 
Those who have studied Balaeniceps s affinities from its skeleton seem to have 
been mainly concerned with its heron-like or stork-like features, and have neglected 
to consider its likeness to the pelicans. Jardine (1852) may have been responsible 
for this when he noted what he thought were significant differences from the " true 
pelicans ". Earlier impressions of Balaeniceps however were that it was near the 
pelicans. For example, its first mention in literature was by Ferdinand Werne 
(1848 : 143) who recorded that on 15th December 1840, " During my siesta someone 
saw a water bird that seemed to be as big as a young camel, which actually had a 
beak just like a pelican's, only without the pouch beneath it ". Even Gould (1852) 
referred to it as a kind of pelican, and Chalmers Mitchell (1913 : 701) considered 
that this opinion was " at least as happy as the more confident statements of later 
writers". None of the non-pelican characters given by Jardine are skeletal. No 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 67 

evidence from comparative osteology has been given as a reason for not putting 
Balaeniceps with the Pelecaniformes, although this has usually been implied on the 
few occasions when differences between Balaeniceps and the Pelecaniformes have 
been described. For example, Parker (i860 : 329) when describing the sternum, 
mentions that the keel extends to the posterior end of the sternum in Balaeniceps, 
as it does in the storks and herons, " whereas in the Pelicans, Gannets and Cormorants 
it scarcely continues beyond the middle of that bone ". 

On the other hand, skeletal characters common to Balaeniceps and the Pelecani- 
formes have often been referred to. Sometimes they have been attributed to conver- 
gence (Parker 1861 : 308), or to the " common inheritance " of the Pelecaniformes and 
Ciconiiformes (Chalmers Mitchell 1913 : 699), but more often they are mentioned 
without comment or even without reference to the fact that they occur in both 
Balaeniceps and the Pelecaniformes. These characters are summarized below. 
They are arranged under three headings, and when a character is mentioned by more 
than one author, or under more than one heading, it is only referred to the first time 
it occurs on the list. Only original works are referred to. An asterisk is placed 
against the characters considered in the present investigation. 



CHARACTERS OF BALAENICEPS'S SKELETON THAT ALSO 
OCCUR IN THE PELECANIFORMES. 

a. Noted and commented on by authors 

Parker (1861 : 308) 

* The palatines have the " same essential structure " in other fish eating 
birds, such as the Pelican, Cormorant and Gannet, because the " motions 
of the upper jaw on the cranium " are the same. 

Chalmers Mitchell (1913 : 699) 

* Long lachrymals. 

* Mesial ankylosis of the palatines. 
Shell-like par occipital processes. 

* Clavicle ankylosed to carina sterni. 
Shape of the head of the humerus. 

He says these are either due to " convergent modifications between birds which, 
after all, are not very far apart in the system " or to the " common inheritance " of 
the Pelecaniformes and " their immediate allies ". 

Bohm (1930 : 700) 

Balaeniceps resembles Pelecanus in its closed palate. 

* Hook to premaxilla. 

* Bony nasal septum. 
Complete interorbital septum. 
Well developed post orbital process. 

Lack of a post angular process on the lower jaw. 



68 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

He says the Pelecaniformes are more like the herons than the storks, except for 
Pelecanus which is atypical, and more like the storks and Balaeniceps. 

B. Noted by authors, without comment 

Parker (i860) 

Cervical vertebrae have haemal arches (p. 328). 

* Furculum articulates with acrocoracoid flange (p. 329). 
Tongue is small (p. 330). 

Parker (1861) 

" General class resemblance " in occipital region (p. 275). 
Sudden bend in furculum (p. 340). 

Chalmers Mitchell (1913) 

* Nasal groove (p. 690). 

c. Mentioned by authors, without reference to the Pelecaniformes 

Parker (1861) 

* Arrangement of the articulating surfaces of the quadrate and lower jaw 
(p. 310). 

Ischium is longer than ilium posteriorly (p. 337) . 

No prepubic process (p. 337). 

Wing skeleton — Parker says this is like that of the herons (p. 342) ; I found 

it as much like that of Pelecanus as Ardea. 

Well developed " cnemial ridges " in tibio-tarsus (p. 343). 

Slight sigmoid curve at distal end of tibio-tarsus (p. 343). 

" Anterior cavity " at proximal end of tarso-metatarsus is deeper than 

in Ardea (p. 343). 

* Complex hypotarsus (p. 344). 

* Mobile hallux (p. 344). 

Chalmers Mitchell (1913) 

A rounded notch separates the metasternum from the posterior lateral 

processes of the sternum (p. 694). 
Bases of coracoids do not meet in the mid-line (p. 696). 
A notch separates the posterior ends of the ilium and ischium (p. 696). 
No horizontal ridge formed by the " dorso-lateral edge of the post-acetabular 

ilium " (p. 696). 
A tibial bridge is present (p. 697). 

Of these skeletal characters common to Balaeniceps and the Pelecaniformes, 
some have been described as being due to " convergence " and others to " common 
inheritance ". It is one of the problems of taxonomy to distinguish between these 
two causes. In this instance the problem is to determine what are significant 
taxonomic characters in the pelecaniform skeleton. At the same time it might not 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 69 

be out of place to refer to the wider problem of the taxonomic significance of 
osteological characters One point of view is that bone is not an easily adapted 
substance, and that therefore phylogeny is readily determined from an examination 
of the skeleton. Verheyen (1953 : 480), for example, says that " systematics 
based on comparative osteology is perfectly realizable " since osteological characters 
are " practically invariable " and are " sheltered from the adaptations and modifica- 
tions imposed by frequent habit ". There is clearly a good deal of truth in this, 
but it is a point of view that should be regarded with caution. There is evidence to 
show that bone is a plastic substance readily moulded by any change in the forces 
exerted by the muscles attached to it. This opinion is expressed, for example, by 
Weinmann & Sicher (1947 : 120) who say, "if it be true that functional stresses 
shape the bone, then it is equally true that a change of strength or direction of 
forces will lead to changes in the form and structure of bones ". Changes in muscle 
function related to changes in habit are therefore reflected in the skeleton. 
Similarities in habit of unrelated species and differences in habit of related species 
can produce a crop of adaptive osteological characters which may obscure phylogeny. 
Phylogeny may be apparent only in a number of small characters which have been 
relatively unaffected by adaptive changes. The sum of these characters may be 
peculiar to a particular group. Although the members of such a group vary in 
appearance and habit, and show convergence with other groups, they will have most 
of the small characters typical of their group. These " non-adaptive " characters 
differ from group to group and may occur in different parts of the skeleton, so that 
each group must be studied separately to get the " feel " of its typical characters. 

In the light of these observations it will be understood that a " pelicaniform 
character " is hard to define precisely. The skeletal characters considered here are 
mainly those which distinguish the Pelecani and Fregatae from the Ciconiidae and 
Ardeae. Some of them, for example the acrocoracoid flange, also occur in other 
groups of birds. For this reason authors have not regarded it as important that 
Balaeniceps has them (Chalmers Mitchell, 1913 : 695). However, they have been 
included here because it is now understood that any given character may be 
taxonomically significant in one group, but not necessarily in another (e.g. see 
Cain, 1954 : 268). The acrocoracoid flange distinguishes the Pelecani, Fregatae and 
Balaeniceps from the Ciconiidae and Ardeae, but not from the Scopidae, 
Falconiformes, most Charadriiformes, Columbidae, Strigiformes, some Procel- 
lariiformes, and many other groups. It is not intended to imply that all the groups 
with an acrocoracoid flange are related, or that any of them are necessarily more 
closely related to the pelicans than to the storks. 

Not all the " pelicaniform characters " considered here occur throughout the 
Pelecani and Fregatae. Sometimes one genus, or more, may differ in one feature 
from the others. For example, adults of Anhinga and Sula have no hook at the tip 
of the premaxilla, but they are typical in most other respects. 

In other cases there may be a general trend, or tendency within the group, towards 
a certain condition, though all the genera are not necessarily concerned in it. One 
example of this is the tendency for reduction of the antorbital vacuity. In 



70 THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 

Pelecanus, the antorbital vacuity is large, as it is in most birds ; in Fregata, Phala- 
crocorax, Anhinga and Sula it becomes progressively reduced. Balaeniceps, there- 
fore, with no antorbital vacuity, would complete the series. 

Sometimes an underlying pattern can be traced in a structure, with differences 
in details in each genus. A good example of this is the arrangement of the quadrate 
condyles in the jaw articulation (see Text-figures). The general form of the nasal 
cavity and of the palate possibly come into this category. In each of these instances 
Balaeniceps has the same underlying pattern as the Pelecaniformes, but the storks 
and herons do not. 

To sum up, the general skeletal features which can be described as " pelecaniform " 
and which occur in Balaeniceps but not in the storks and herons are as follows : 

(i) Position of nasal groove along upper mandible, and strong terminal 
hook (see a(i) and a(2)). 

(2) Arrangment of nasal cavity (see A (3) and A (4)). 

(3) Relationship of bones of palate and maxillopalatines (see a(5)). 

(4) Size of lachrymal and antorbital vacuity (see A (6)). 

(5) Type of jaw articulation (see a(j)). 

(6) Some features of the pectoral girdle and sternum (see b) . 

(7) Shape of first metacarpal (see d(3)). 

In my opinion the skeleton of Balaeniceps has many points of similarity, due to 
convergence, with the Ciconiidae and Ardeae, but, in spite of its difference in out- 
ward appearance from any of the Pelecaniforms, it shares several apparently non- 
adaptive features with them. I find it difficult to account for this unless Balaeniceps 
is more closely related to the Pelecaniformes than it is usually considered to be. 
Therefore, from a consideration of the skeletal characters of Balaeniceps rex, it seems 
that this species could occupy a monotypic family in the order Pelecaniformes, 
possibly near the Pelecanidae. 

SUMMARY 

1. A number of features of the skeleton of Balaeniceps rex were found to be more 
like the pelicans than either the storks or herons, with which Balaeniceps is usually 
grouped. 

2. A study of the literature showed that the pelican-like characters of Balaeniceps 
had never been fully investigated. 

3. The skeleton of Balaeniceps was compared with those of all the families of 
the Pelecaniformes, except the Phaethontidae, and with the Ardeidae, Cochlearidae 
and Ciconiidae of the Ciconiiformes. Reasons are given for limiting comparison 
to these groups. 

4. The characters common to Balaeniceps and the Pelecaniformes are described 
in detail. 

5. The osteological evidence suggests that Balaeniceps is more closely related to 
the Pelecaniformes than to the Ciconiiformes, and the family Balaenicipitidae may 
reasonably be placed in the Pelecaniformes, possibly near the Pelecanidae. 



THE PELECANIFORM CHARACTERS OF THE SHOE-BILL STORK 71 

REFERENCES 
Bartlett, A. D. 1861. On the affinities of Balaeniceps. Proc. Zool. Soc. London, 1861 : 

I3I-I34- 

Beddard, F. E. 1888. On certain points in the visceral anatomy of Balaeniceps rex bearing 

on its affinities. Ibid., 1888 : 284-290. 
Bohm, M. 1930. tlber den Bau des jugendlichen Schadels von Balaeniceps rex nebst Bemer- 

kungen iiber dessen systematische Stellung und fiber das Gaumenskelett der Vogel. Z. 

Morph. Okol. Tiere, Berlin, 17 : 677-718. 
Bonaparte, C. L. J. L. 1855. Conspectus generum avium, 1849-57. Lugduni Batavorum. 
Cain, A. J. 1954. Subdivisions of the genus Ptilonopus. Bull. Brit. Mus. (nat. hist) Zool. 

2 (8) : 267-284. 
Des Murs, M. O. 1859. Considerations oologiques sur l'oiseau type du genre Baleniceps. 

Rev. et Mag. Zool. Ser. 2, 11 : 477-481. 
Furbringer, M., 1 888 Untersuchungen zur Morphologic und Systematik der Vogel. 2 : 1 187-1 196. 

Amsterdam. 
Gadow, H. 1893. Vogel : Aves in Bronn's Klass. n. Ord. des. Thier-Reichs . Bd. 6, Abth 4. 

Leipzig. 
Giebel, C. G. 1873. Balaeniceps rex. Z. ges. Naturw. Berlin, 41 : 350-354. 
Glenny, F. H. 1955. Modifications of pattern in the aortic arch system of birds and their 

phylogenetic significance. Proc. U.S. nat. Mus. No. 3346, 104 : 525-621. 
Gould, J. 1852. On a new and most remarkable form in ornithology. Proc. Zool. Soc. 

London, 1851 (published 1852) : 1-2. 
Heuglin, M. T. von. 1856. Systematische Uebersicht der Vogel Nord-Ost-Afrika's. Wien. 

■ 1873. Ornithologie Nordost-Afrika's, 3 : 1095-1099. Cassel. 

Jardine, W. 1852. Ornithology in 1850 ; Balaeniceps rex. Contr. Orn. 1851 (published 1852) : 

11-14. 
Mayr, E. & Amadon, D. 1951. A classification of recent birds. Amer. Mus. Novit. 1496. 
Mitchell, P. C. 1913. Observations on the anatomy of the Shoe-bill [Balaeniceps rex) and 

allied birds. Proc. Zool. Soc. London, 191 3 : 644-703. 
Parker, W. K. i860. Abstract of notes on the osteology of Balaeniceps rex. Ibid., i860 : 

324-330. 
1861. On the osteology of Balaeniceps rex. Trans. Zool. Soc. London, 4 : 269-352. 

1862. [Letter on Reinhardt's 1861 paper.] Ibis, London : 297-299. 

Pycraft, W. P. 1898. Contributions to the osteology of birds. I. — Steganopodes. Proc. 

Zool. Soc. London, 1898 : 83. 
Reinhardt, J. i860. On the affinities of Balaeniceps. Ibid., i860 : 377-380. 

1861. Some remarks on the genus Balaeniceps. Ibis, London, 1862 : 158-175. [Trans- 
lation by A. Newton of article in K. danske Vidensk. Selsk. Kobenh. 1861 : 135-154.] 

Shufeldt, R. W. 1901. Notes on the osteology of Scopus umbretta and Balaeniceps rex. 

J. Anat. Physiol. 2 : 405-412. 
Stresemann, E. 1927-34. Aves in Kukenthal & Krumbach's Handb. d. Zool. Bd. 7 Hft. 2. 

Berlin and Leipzig. 
Technau, G. 1936. Die Nasendriise der Vogel. /. Orn. Leipzig. 84 : 511-617. 
Verheyen, R. 1953. Bijdrage tot de osteologie en de systematick der Anseriformes. Gerfaut, 

Bruxelles, 43. Supplement : 373-500. 
Weinmann, J. P. & Sicher, H. 1947. Bone and Bones. London. 

Werne, F. 1848. Expedition zur Entdeckung der Quellen des Weissen Nil (1840-41). Berlin. 
Wetmore, A. 1930. A systematic classification for the birds of the world. Proc. U.S. nat. 

Mus. 76, 24 : 1-8. 

1951. A revised classification for the birds of the world. Smithson. misc. Coll. 117, 

4 : 1-22. 



PLATE 
[Scale : The skulls have been variously reduced, so that the crania are of approximately 
the same size. The actual total length of each skull is given below in brackets.] 
Lateral views of skulls of : 



(i) Balaeniceps rex (265 mm.). 

(2) Pelecanus onocrotalus (410 mm.). 

(3) Sula bassanus (180 mm.). 

e = external naris. 

g = nasal groove. 

h = premaxillary hook. 

1 = position of internal nares. 



(4) Ciconia ciconia (255 mm.). 

(5) Ardea goliath (235 mm.). 

(6) Cochlearius cochlearius (125 mm.) 

l = lachrymal. 

m =maxillopalatine. 

p = palatine. 

v = antorbital vacuity. 




Bull. B.M. (N.H.) Zool. 5, 3. 



PLATE 3 







PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 
BARTHOLOMEW PRESS, DORKING 



4^ « 

.' 6 CO 

A REVISION OF THE 

LAKE VICTORIA HAPLOCHROMIS 

SPECIES (PISCES, CICHLIDAE) 

PART II: H. S,4l7K,4GEJ (PFEFFER), 
H. PRODROMUS TREWAVAS, 

H. GRANTI BLGR., 
AND H. XENOGNATHUS SP. N. 



P. H. GREENWOOD 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 4 

LONDON: 1957 



A REVISION OF THE LAKE VICTORIA 

HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) 

PART II: H. SAUVAGEI (PFEFFER), 

H. PRODROMUS TREWAVAS, 

H. GRANTI BLGR., 
AND H. XENOGNATHUS, SP.N. 



BY 

P. H. GREENWOOD 

East African Fisheries Research Organization, Jinja, Uganda. 



Pp. 76-97 ; Plate 4 ; 8 Text-figs. 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 4 

LONDON: 1957 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

This paper is Vol. 5, No. 4 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued October, 1957 Price Eight Shillings 



A REVISION OF THE LAKE VICTORIA 
HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) 

PART II 1 : H. SAUVAGEI (PFEFFER), 

H. PRODROMUS TREWAVAS, 

H. GRANT I BLGR. 

AND H. XENOGNATHUS, SP. N. 

By P. H. GREENWOOD 

CONTENTS 

Page 
Introduction ........... 76 

Haplochromis sauvagei (Pfeffer) ....... 76 

Synonymy and description . . . . . . 77 

Distribution .......... 80 

Ecology .......... 80 

Diagnosis and affinities . . . . . . . .80 

Study material and distribution records . . . . .81 

Haplochromis prodromus Trewavas . . . . . . .82 

Synonymy and description . . . . . . .82 

Distribution .......... 85 

Ecology .......... 85 

Study material and distribution records ..... 85 

Diagnosis and affinities . . . . . . . .86 

Haplochromis granti Boulenger ....... 86 

Synonymy and description . . . . . . .86 

Distribution .......... 89 

Ecology ........... 89 

Diagnosis and affinities ........ 89 

Study material and distribution records ..... 90 

Haplochromis xenognathus sp. nov. ....... 90 

Synonymy and description . . . . . . • 9 1 

Distribution .......... 94 

Ecology .......... 94 

Diagnosis and affinities ........ 94 

Study material and distribution records ..... 95 

Discussion ........... 95 

Summary ......... 97 

Acknowledgments .......... 97 

References ........... 97 

1 Part I, see Greenwood, 19566. 

zool. 5, 4. 4§ 



76 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

INTRODUCTION 

Morphologically, the H. sauvagei complex stands apart from any other species- 
group in Lake Victoria. The principal group character is that of the dentition 
which combines recurved outer teeth with multiseriate inner tooth-bands (Text- 

fig- 3). 

Furthermore, the shape of the neurocranium, although differing intra-specifically 
within the group, is unlike that of other Haplochromis. This character is probably 
associated with the multiseriate dentition and relatively powerful jaw musculature. 
Indeed, amongst the non-piscivorous predators such marked divergence in cranial 
anatomy is otherwise only found in mollusc-eating species with hyper-developed 
pharyngeal bones and musculature. Two specifically constant forms of neurocranium 
occur in the " sauvagei " group, but neither can be correlated with the type of 
dental pattern present. 

Trophically, members of the group may be classed as mollusc eaters, although 
available data indicate that other food organisms do contribute to their diet, 
usually in a subsidiary capacity. Unlike other mollusc-eating Haplochromis in this 
lake, species of the " sauvagei " group do not swallow the shells of their prey, but 
remove the soft parts before ingestion takes place. In this respect the feeding method 
is like that of Macropleurodus bicolor (Blgr.), a monotypic genus apparently derived 
from this group. 

Haplochromis sauvagei (Pfeffer), 1896 
(Text-fig. 1 and PL 1 upper fig.) 

Ctenochromis sauvagei Pfeffer, 1896, Thier. Afr. Fische, 15. 

Haplochromis nuchisquamulatus (part), Boulenger, 1915, Cat. Afr. Fish., 3, 290. 

Paratilapia granti (part), Boulenger, 1915, op. cit., 342. 

Paratilapia bicolor (part) Boulenger, 1915, op. cit., 346. 

Paratilapia retrodens (part), Boulenger, 191 5, op. cit., 235. 

Haplochromis sauvagei (part), Regan, 1922, Proc. zool. Soc, Lond., 167. 

? Paratilapia crassilabris (part), Boulenger, 191 5, op. cit., 345. 

I was unable to examine the holotype of H. sauvagei which was mislaid during 
the 1939-45 war ; at present the Berlin Museum authorities cannot confirm whether 
this specimen has been lost. Pending more definite information, no neotype can be 
selected, but, should such a step become necessary, I suggest that the specimen 
B.M. (N.H.) Reg. No. 1956. 9. 17. 1, a male from Entebbe (Text-fig. 1) be given 
neotypical status. 

Fortunately, Pfeffer's original description of Ctenochromis sauvagei is compre- 
hensive, and, when coupled with a photograph of the type, clearly indicates to which 
Haplochromis species his specimen should be referred. The photograph, preserved 
in the British Museum (Natural History), is reproduced in Plate 1. 

Additional material discloses only one important discrepancy with the original 
description, in which the mouth and lower- jaw profile are described as rising steeply : 

. . . ; das untere Profil der Unterkinnlade steigt viel starker. Die von dicken 
und breiten Lippen umgebene kurze Mundspalte steigt nach vom sehr steil auf." 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



77 



In most specimens the ventral head profile is almost horizontal, or, at most, slightly 
oblique. A possible explanation for this discrepancy may lie in the fact that Pfeffer's 
description was taken from a fish preserved with its mouth open. From the photo- 
graph it is clear that, if the jaws were restored to their natural position, the cleft 
and lower jaw profile would be slightly oblique. 

The present synonomy for H. sauvagei is essentially that prepared by Regan 
(1922), but some of the specimens which he referred to this species are now placed 
in others. In this connection, reference should be made to the list of study material. 

Paratilapia crassilabris part (Boulenger, 1915) is tentatively retained in the 
synonomy on the basis of a single specimen (B.M. (N.H.) Reg. no. 191 1.3. 3. 32). 
This individual cannot be identified with certainty, but it is nearer H. sauvagei than 
any other species with thickened lips and dentition not of the generalized type. 



Fig. 1. Haplochromis sauvagei, g, B.M. (N.H.) 1956.9. 1 7.1. Drawn by Miss L. Buswell. 



Description. Based on 85 specimens, 58-105 mm. standard length. Of the 
measurements made, only cheek depth clearly shows allometry with standard 
length. 

Depth of body 30-4-41-8, mean (M) = 35-6 ; length of head 29-6-34-5 (M = 31-9) 
per cent of standard length. Dorsal head profile varying from decurved to straight, 
but strongly sloping, the former shape occurring more frequently. 

Preorbital depth 15-4-20-2 (M = 17-3) per cent of head length, least interorbital 
width 23-0-31-2 (M = 27-0) per cent. Snout as broad as long, its length 27-2-35-5 
(M = 30-8) per cent of head ; eye diameter 25-7-33-4 (M = 28-9) per cent. Cheek 
becoming relatively deeper with increasing standard length ; four size-groups are 
recognized, 58-69 mm. S.L. (N = 13), 70-80 (N = 21), 81-90 (N = 27) and 91- 
105 (N = 24), for which the cheek depth is 21-0-25-0 (M = 23-2), 20-4-26-0 (M 
= 23-7), 22-0-26-9 (M = 24-4) and 24-1-26-6 (M = 25-1). 



78 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Caudal peduncle I3'9~i9*3 (M = 16-4) per cent of standard length ; its length 
i-i-i-6, times its depth. 

Mouth horizontal or slightly oblique ; posterior maxillary tip reaching or almost 
reaching the vertical to the anterior orbital margin. Lips thickened ; the depth of 
the upper lip, measured mid-laterally, is contained 4-4J times in the eye-diameter. 
Jaws equal anteriorly, the lower 30-6-37-7 (M = 34-5) per cent of head length and 
1-0-1-5 (mode 1-3) times as long as broad. 

Gill rakers short, 7-9 (rarely 10) on the lower limb of the first arch. 

Scales ctenoid ; lateral line with 31 (f.7), 32 (f.29), 33 (f.39), 34 (f.9) or 35 (f.i) 
scales ; cheek with 3-4 (rarely 2) series. 7-9 (less frequently 6) scales between dorsal 
fin origin and upper lateral line ; 7 or 8 (less commonly 6 or 9) between pectoral 
and pelvic fin-bases. 

Fins. Dorsal with 24 (f.n), 25 (f.56) or 26 (f.18) rays, anal 10 (f.i), n (f.io), 12 
(f.71) or 13 (f.3), comprising XV-XVII, 8-10, and III, 7-10 spinous and branched 
rays for the fins respectively. Pectoral slightly shorter than the head, or occasionally 
of equal length. Pelvic with the first ray produced, variable in its posterior extension 
but longer in adult males than females. Caudal sub-truncate. 

Lower pharyngeal bone triangular, its dentigerous area i^-ij times as broad as 
long ; pharyngeal teeth slender and cuspidate. In some specimens, teeth in the 
median rows are slightly enlarged, but retain their bicuspid crowns. 

Teeth. In the outermost series of both jaws, the teeth have strongly recurved tips 
and are unequally bicuspid or unicuspid. The predominant tooth form is apparently 
correlated with length. Fishes less than 80 mm. S.L. have mainly bicuspid teeth, 
those in the range 80-90 mm. have either unicuspids or an admixture of uni- and 
bicuspids, whilst larger individuals possess mainly unicuspid teeth. When both 
types of teeth are present, the unicuspid form usually occurs anteriorly and laterally. 
There are 32-56 (mode 42) outer teeth in the upper jaw. 

Inner teeth are either tri- or unicuspid ; as in the outer series, unicuspid teeth 
are commoner in fishes above 80 mm. S.L. Antero-medially, the teeth are arranged 
in a broad band comprising 3-8 (mode 4) and 2-6 (modes 3 and 4) rows in the upper 
and lower jaws respectively. Laterally, the band narrows to a single series. A 
distinct inter-space separates the inner and outer series. 

Syncranium and associated musculature. Neurocranial form in H. sauvagei departs 
quite considerably from the generalized Haplochromis type, and approaches that 
of Macropleurodus bicolor (Greenwood, 1956a). Essentially the same points of dif- 
ference with the generalized type occur in both species. The skull has a fore- 
shortened appearance due to the strongly decurved and almost vertically disposed 
ethmo vomerine region. This curvature affects the morphology of the entire pre- 
orbital skull which is less gently curved than in the generalized neurocranium. 

On the other hand, the jaws do not exhibit such radical departure from the 
generalized condition. The premaxilla, apart from a slight broadening of its denti- 
gerous area, compares closely with that of other Haplochromis ; the dentary is 
somewhat shorter, more massive and has a wider median dentigerous area than is 
common in generalized species. Consequent upon these modifications slight differ- 
ences are apparent in the suspensorium. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 79 

Muscle disposition and form are similar to those of basic Haplochromis species. 
However, the adductor mandibulae I is slightly longer (38-44 per cent head length 
compared with 36-39 per cent) and broader (length /breadth ratio 3-0-37 cf. 

4-4-5-5). 

The syncranium, its musculature and the dentition all foreshadow the condition 
found in H. prodromus, and therefore that which reaches its ultimate expression in 
the genus Macropleurodus Regan (Greenwood, op. cit.). It is perhaps significant 
that variability in the degree to which the syncranium departs from the basic type 
is greater in H. sauvagei than in H. prodromus. 

Causal factors responsible for the characteristic preorbital face in both H. sauvagei 
and H. prodromus are not readily determined. From an examination of larval fishes 
it is manifest that, as in M. bicolor, this form develops during post-larval ontogeny. 
It is probably effected by differential growth of various syncranial parts, especially 
since the ethmovomerine region is not directly affected by the moulding influence of 
muscle insertions. However, the premaxilla, which is closely associated both anato- 
mically and functionally with the dentary, could exert considerable influence over 
this region. In H. sauvagei the dentary is short in relation to the head and also in 
comparison with other Haplochromis species of comparable size. If, during post- 
larval ontogeny, this bone increased in length more slowly than the neurocranium, 
and if it is to remain functionally integrated with the upper jaw, then there can be 
two morphological results : either a skull of the H. sauvagei type, or one in which 
the upper jaw projects anteriorly beyond the lower. A third possibility, that the sus- 
pensorium be rotated anteriorly, cannot be considered in this case, since its almost 
vertical alignment in H. sauvagei is typically that of the basic type. 

H. sauvagei includes Gastropoda as a substantial part of its diet. As in H. prodromus 
and M. bicolor the soft parts alone are ingested. The feeding habits of M. bicolor 
have been described elsewhere (Greenwood op. cit.) : when feeding on snails, 
aquarium-kept H. sauvagei follow the same general pattern, except that after 
grasping the foot of the snail between its jaws the fish then uses the shell as a fulcrum 
to lever out the soft parts. Only rarely is the shell crushed by the jaws. 

Coloration in life : Breeding males. Ground colour dark grey-green or blue-grey, 
lighter or yellowish ventrally ; a suffused coppery sheen on the flanks and ventral 
aspects of the operculum. Dorsal fin black basally, becoming slate-coloured distally ; 
lappets red ; red spots, often coalesced, between the soft rays. Anal dark with a 
red flush ; ocelli yellow. Caudal dark grey proximally, lighter distally, and with 
an overall orange-red flush. Pelvics black laterally, orange-red medially. Non- 
breeding males have similar coloration except that the copper flush is absent and 
other bright colours are less intense. Females and immature males. Ground col- 
our golden-green, shading to pearly- white ventrally. All fins yellow-green. 

In both sexes there may develop after death a dark longitudinal band running 
mid-laterally from the eye to the dorsal fin base, a second band running dorso- 
laterally approximately along the upper lateral-line, and 6-10 narrow transverse 
bars across the flanks. In life these markings are rarely discernible. 

Amongst females a second type of coloration is known. This takes the form of 
irregular black blotches on a yellow ground and is identical with the bicolor pattern 

zool. 5, 4. 4§§ 



80 REVISION OF LAKE VICTORIA H APLOCHROMIS SPECIES 

described for certain female Macropleurodus bicolor, Hoplotilapia retrodens, and 
Haplochromis nigricans (Greenwood, 1956*2 and b). 

Since collectors show some predilection for fishes with a striking colour pattern, 
it is difficult to obtain accurate frequency estimates for the bicolor pattern. In the 
present sample, 25 per cent of females are bicolor. As most specimens were obtained 
by collectors aware of possible biasing factors, this figure may be accepted as fairly 
reliable. No male bicolor variants have yet been recorded. Thus, the incidence of 
bicolor variants seems sufficiently high to recognize the phenomenon as sex-limited 
polychromatism, and not merely the maintenance of an atypical genotype by re- 
current mutation. Aberrantly coloured females were found in most localities. None 
exhibits a pattern intergrading with that usual for females. 

Sex-limited polychromatism involving the same phenotypic expression was 
observed in M. bicolor and Hoplotilapia retrodens (Greenwood, 1956a). It seems 
probable that hypotheses regarding its genie basis and evolutionary significance in 
these species are also applicable to H. sauvagei. The possible significance of bicolor 
females as indicating phyletic relationship amongst the various species in which 
they occur has also been discussed (Greenwood, op. cit.). It was concluded that, in 
general, no reliability could be placed on this character, and that its repeated appear- 
ance was probably attributable to the oligophyletic origin of the Lake Victoria 
species-flock. Nevertheless, it is suggestive that both H. sauvagei and M. bicolor 
exhibit " bicolor " polychromatism as well as an apparent similarity in fundamental 
syncranial morphology. 

Colour in preserved material : Adult males. Slate-grey to sooty, the longitudinal 
and transverse banding often obscured. Spinous dorsal fin grey, soft part hyaline 
but maculate. Anal and caudal hyaline. Pel vies black on the outer half, hyaline 
mesially. A dark lachrymal stripe and two bars across the snout are often present. 
Females and immature males. Ground colour variable, from silver-grey to brownish. 
Banding, as described above, usually developed. All fins hyaline, the soft dorsal 
and upper half of the caudal, maculate. 

Distribution. Known only from Lake Victoria. 

Ecology : Habitat. Restricted to littoral zones where the bottom is hard (sand or 
shingle) ; the species is especially common over exposed sandy beaches. 

Food. The gut contents of forty-five fishes from various localities indicate that 
H. sauvagei feed mainly on Gastropoda (f.19), bottom deposits, which included insect 
larvae, Copepoda and diatoms (f.19), and Insecta (chiefly larval boring may-flies, 
Povilla adusta Navas) (f.4). No fragments of snail shell were observed, although 
opercula occurred frequently in the stomach and intestine (see also p. 79). 

Breeding. Spawning sites and behaviour are unknown. In many localities, sexually 
active and quiescent fishes, and brooding females occur together. 

The smallest adult fish was a female 72 mm. S.L. All specimens over 80 mm. 
were adult. No difference was detected in the sizes of adult males and females. 

Diagnosis. H. sauvagei is distinguished from other Haplochromis in Lake Victoria 
by combinations of the following characters : lips thickened ; outer teeth with 
strongly recurved tips ; usually more than three inner rows of teeth in the upper 
jaw (mode 4). The species closely resembles H. prodromus, from which it may be 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 81 

separated by its slightly thinner lips and smaller adult size. In life, male breeding 
coloration serves to separate the two species. 

Affinities. Similarity in the skull architecture and the dentition of H. sauvagei 
and H. prodromus suggest a phyletic relationship between the species. Consequent 
upon these anatomical similarities, the species show a close parallel in their feeding 
habits and food preferences, although in this respect H. sauvagei may be considered 
less specialized than H. prodromus. 



Study material and distribution records 



Museum and Reg. No. 



Uganda 



B.M. (N.H.) 1908.5.30.365-366 (as Paratilapia 

granti) 
B.M. (N.H.) 1906. 5. 30. 371-372 (as P. granti) 
B.M. (N.H.) 1906. 5. 30. 413 
1911. 3-3-27 • 
1909.3.29.9 - 
(all as P. bicolor) 
B.M. (N.H.) 1909. 5. 11. 11 

„ 1906.5.30.374-377 

1956.9. 17. 1 (See Text-fig. 1 
1956. 10.9. 1-25 

„ „ 26-30 . 
„ „ 31-34 • 

„ „ 35-36, 201 
„ „ 37-40 . 



41 
42 

43 



Locality. 


Collector 


Bunjako 


. Degen. 


Bugonga (Entebbe) 
Sesse Is. 


. Bayon. 



Bunjako 

Entebbe, Airport beach 

Entebbe, harbour 
Bugungu (Napoleon 

Gulf) 
Jinja pier 
Shore opposite Kirinya 

Point (Napoleon 

Gulf) 
Kirinya Point 
Old Bukakata 
Katebo 



44 

45-72 

73 

74-75 



76 

77-80 

81 
82 



Tanganyika Territory 

Mwanza 
. Majita 

Ukerewe Is. 
Bukoba 



Kenya 



83 



Kisumu 

Kamaringa (Kavirondo 

Gulf) 
Kach Bav (Kavirondo 

Gulf) 
Open water 5 miles N. 

of Kendu (Kavirondo 

Gulf) 
Rusinga Island 



Degen. 
E.A.F.R.O. 



82 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Haplochromis prodromus Trewavas, 1935 
(Text-figs. 2 and 3) 

Paratilapia retrodens (part), Boulenger, 1915, Cat. Afr. Fish., 3, 235. 
Haplochromis ishmaeli (part), Boulenger, 191 5, op. cit., 293. 

Haplochromis annectens Regan 1922 (nee. Cyrtocara annectens Regan, 1921), Proc. zool. Soc. 
Lond., 167, fig. 2. 

Description. Based on sixty-two specimens (including the holotype), 68-130 mm. 
S.L. None of the morphometric characters studied shows allometry with standard 
length. 

In its general appearance H. prodromus closely resembles H . sauvagei, from which 
species it is distinguished by its thicker lips, slightly deeper cheek and larger adult 
size. 




Fig. 2. Haplochromis prodromus, g, holotype (from Regan, the cichlid fishes of Lake Victoria, 

Proc. Zool. Soc, 1922, 168, fig. 2). 

Depth of body 32-8-40-0 (M =36.2) ; length of head 29-4-33-6 (M = 31-5) per 
cent of standard length. Dorsal head profile somewhat variable, but always curved ; 
strongly decurved in some large individuals, less so in smaller fishes (70-75 mm. S.L.). 

Preorbital depth 14-0-19-1 (M = 15-8) per cent head length ; least interorbital 
width 24-0-31-3 (M = 28-1) per cent. Snout as broad as or slightly broader than 
long, rarely longer than broad, its length 27-5-36-8 (M = 32-7) per cent of head ; 
eye diameter 25-8-33.3 (M = 27-8) ; cheek 22-0-30-5 (M = 26-7) per cent. 

Caudal peduncle 12-6-18-1 per cent of standard length, its length 1-0-1-7 ( m °de 
1-3) times its depth. 

Mouth horizontal ; posterior maxillary tip reaching or almost reaching the vertical 
to the anterior orbital margin. Lips thickened ; the depth of the upper lip, measured 
mid-laterally, contained 3-3^ times in eye diameter. Jaws equal anteriorly, or 
infrequently the lower very slightly shorter ; lower jaw 30-5-37-8 (M = 34-3) per 
cent of head length, up to 1-3 (mode i-i) times as long as broad. 

Gill rakers short, 7-9 on the lower limb of the anterior arch. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 83 

Scales ctenoid; lateral line with 30 (f.i), 31 (f.7), 32 (f.16), 33 (f.35) or 34 (f.2) 
scales ; cheek with 3 or 4 series. 7 or 8 (rarely 6J or 9) scales between origin of 
dorsal fin and the lateral line, 7 or 8 (less frequently 9) between pectoral and pelvic 
fin bases. 

Fins. Dorsal with 24 (f.7), 25 (f.40) or 26 (f.15) rays, anal with n (f.7), 12 (f.47) 
or 13 (f.8), comprising XV-XVII, 8-10 and III, 8-10 spinous and branched rays 
for the fins respectively. Pectoral shorter than the head. Pelvic fins with the first 
ray produced and of variable posterior extension, but reaching the anal fin in most 
adult fishes. Caudal sub-truncate. 

Lower pharyngeal bone triangular, its dentigerous surface about ij times as broad 
as long ; pharyngeal teeth slender and cuspidate ; those of the median series some- 
times enlarged. 

Teeth. The dental pattern and tooth form in H. prodromus closely resemble those 
of H. sauvagei. 

In the outer series of both jaws the teeth have strongly recurved tips and are 
unequally bicuspid or unicuspid. Bicuspid and weakly bicuspid teeth are the pre- 
dominating forms in fishes less than 100 mm. S.L. Above this size most teeth are 
unicuspid. 26-56 (mode 40) outer teeth occur in the upper jaw. 

Inner teeth are either tri- or unicuspid, the tricuspid form occurring most fre- 
quently in fishes less than 90 mm. S.L. Antero-medially the teeth are arranged in 
3-7 (modes 4 and 5) and 3-6 (modes 3 and 4) series in the upper and lower jaws 
respectively. The posterior medial margin of the upper tooth-band is straight or 
slightly curved, that of the lower band is distinctly curved (Text-fig. 3). 

The dental pattern of the holotype must be considered aberrant ; it is not re- 
peated in any of the sixty-one additional specimens. In the type, some postero- 
lateral inner teeth are displaced medially from their series, thereby giving a spurious 
impression of a tooth band widened at that point. There is no increase in the width 
of the underlying premaxillary alveolar surface, nor is there an increase in the 
number of tooth rows (see fig. 14 in Regan, 1922). In all other respects the dentition 
of this specimen agrees closely with those described above. 

Syncranium and associated musculature. The neurocranium and premaxilla of 
H. prodromus are virtually identical with those of H. sauvagei. The dentary, however, 
is relatively more massive and the mental profile is almost vertical. 

Likewise, the jaw musculature compares closely with that of H. sauvagei, except 
that the adductor mandibulae I is somewhat shorter (36-39 per cent head length). 

Observations made on the feeding methods of H. prodromus kept in aquaria, 
indicate that snails are removed from their shells in a manner similar to that 
employed by Macropleurodus bicolor. That is, the shell is crushed free by the jaws 
before ingestion takes place. The species was not seen to lever out the soft parts as 
is usual with H. sauvagei. 

Coloration in life : Adult males. Ground colour slatey blue-grey ; a peacock-blue 
sheen on the belly and ventral flanks. Chest and branchiostegal membrane black, 
operculum with a golden flush. Very faint indications of a dark mid-lateral stripe 
and seven transverse bars ; also a faint lachrymal stripe. Dorsal dark, with a deep 
red flush between both spinous and soft rays ; lappets orange. Anal sooty, ocelli 



84 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



deep yellow. Caudal sooty, with a faint orange flush along its posterior margin. 
Pelvics black. Females and immature males. Ground colour silver-grey above the 
mid-lateral stripe and silver below, with a faint peacock-blue flush on the flanks. 
Transverse barring is indistinct. Dorsal fin dark. Caudal and anal hyaline. Pelvics 
faintly yellow. 





<a 



Fig. 3. 



I cm. 

The premaxillary and mandibular tooth bands in H. pvodromus. 



Colour in preserved material : Adult males. Ground colour dark grey ; in some, 
faint traces of transverse and longitudinal banding. Chest and branchiostegal mem- 
brane black. Dorsal, caudal and anal dark, the soft dorsal maculate. Pelvics black. 
Females and immature males. Pale, banding variable but usually a distinct mid- 
lateral stripe and a faint, more dorsal band running along the upper lateral line ; 
five to nine transverse bars across the flank. All fins hyaline. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



85 



Distribution. Known only from Lake Victoria. 

Ecology : Habitat. Restricted to littoral zones, particularly where the substrate 
is hard (sand or shingle) and occurring less frequently over mud. Thus, the habitat 
of H. prodromus broadly overlaps that of H. sauvagei. Nevertheless, although 
biasing factors are introduced by the size selectivity of sampling gear and the 
limitations imposed by the habitat on the use of certain gear, it seems that H. 
sauvagei occur most frequently over shallow exposed beaches — where H. prodromus 
are less common — and that H. prodromus are more abundant in off-shore to deeper 
waters. This assumption is supported by results obtained when such non-selective 
collecting methods as explosives were used in both habitats. 



Study material and distribution records 
Museum and Reg. No. 



Locality. 



Uganda 



B.M. (N.H.) 1907.5.7.78 (holotype H. prodromus) Buddu coast 
,, 1906.5.30.379 (as P. retrodens) . Bunjako 



B.M. (N.H.) 1956. 10.9 84-97 
„ „ „ 98-99 



100-105 

106 

107 

108-125 

126 

127-129 



130 



Jin] a 

Shore opposite Kirinya 

Point (Napoleon 

Gulf) 
Beach near Nasu Point 

(Buvuma Channel) 
Pilkington Bay 
Hannington Bay 
Entebbe harbour 
Katebo 
Busungwe Bay (Kagera 

river mouth) 
Dagusi Island 



Collector. 



Simon. 
Degen. 
E.A.F.R.O. 



131-133 
134-135 
197-199 



Tanganyika Territory 

Mwanza, Capri Bay 
Godziba Island 
. Majita 



136-137, 196 
138 



Kenya 



Kamaringa (Kaviron- 

do Gulf) 
Kisumu 



Food. Stomach and intestinal contents of seventy-four fishes were examined. Of 
these, eleven were empty, fifty-seven contained only the remains of Gastropoda, 
three contained Gastropoda and Insecta, and three yielded unidentifiable sludge. 
Due to their very fragmentary nature the specific identification of molluscan remains 
was difficult ; where identification was possible the genus Bellamya predominated. 
As many as twenty-two snail opercula were recorded from the intestine of a single 
fish, although the modal estimated number of snails per individual was about four. 



86 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Breeding. Sexually active and quiescent individuals were associated in all localities, 
but no data were collected on breeding sites or spawning behaviour. Only one female 
was found carrying larvae in the buccal cavity. There is apparently no sex-correlated 
adult size difference in this species ; the smallest sexually active individual was a 
male 102 mm. long. 

Diagnosis. The same character complex serves to separate H. sauvagei and H. 
prodromus from the other Haplochromis of Lake Victoria. H. prodromus is distin- 
guished from H. sauvagei by its larger adult size, thicker lips, slightly deeper cheek, 
and, in life, by male breeding coloration. 

Affinities. The apparent phyletic relationship between H. prodromus and H. 
sauvagei on the one hand, and the more specialized Macropleurodus bicolor on the 
other, has been discussed above and elsewhere (Greenwood 1956a). In the latter 
paper, it was shown that Regan's suggested relationship between H. prodromus and 
Platytaeniodus degeni Blgr. can no longer be considered valid. Regan's views were 
based on the type and then unique specimen of H. prodromus whose dental pattern 
is aberrant. In any case, the posterior widening of the premaxillary dental surface 
is apparent and not actual in this fish, whereas in P. degeni the premaxilla undergoes 
a localized but distinct broadening during post-larval ontogeny. 



Haplochromis granti Boulenger, 1906 
(Text-figs. 4 and 5) 

Paratilapia granti (part), Boulenger, 1915, Cat. Afr. Fish., 3, 342, Fig. 231. 
Haplochromis sauvagei (part), Regan, 1922, Proc. zool. Soc, Lond., 167. 

In Regan's revision of the Lake Victoria Cichlidae {ibid., 1922), H. granti was 
treated as a synonym of H. sauvagei. After comparing the type with other specimens 
now available, I conclude that the two species should be regarded as distinct. This 
conclusion is supported by field observations. Both species have in common the 
" sauvagei " group characters of broad inner tooth bands, outer teeth with strongly 
recurved tips, and thickened lips. But they differ considerably in gross morphology 
and in certain details of dental pattern. The holotype of H. granti (figured in 
Boulenger, 1915) does not present a specifically typical appearance. However, its 
dental pattern indicates conspecificity with the specimens here described as H. granti. 
Furthermore, in the type, characters which contribute to gross morphology, for 
instance the form of the dentary and the head shape, intergrade with those of other 
specimens possessing a more typical facies. 

One rather damaged specimen (B.M. (N.H.) Reg. No. 1911.3.3.28), identified by 
Boulenger as Paratilapia retrodens and later by Regan as H. sauvagei, should probably 
be referred to H. granti. Because of this uncertainty P. retrodens is not included in 
the revised synonomy of H. granti. 

Description. — Based on the type, two paratypes and twenty-six additional speci- 
mens in the size range 70-122 mm. S.L. No clear-cut allometry with standard length 
was observed in the morphometric characters listed below. 

Depth of body 327-39*3 (M = 35*4) i length of head 28-8-33-3 (M = 31-5) per 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 87 

cent of standard length. Dorsal head profile slightly curved, or, less frequently, 
straight and gently to steeply sloping. 

Preorbital depth 15-3-19-0 (13-3 in the smallest specimen) (M = 17-1) per cent of 
head length ; least interorbital width 25-0-32-8 (M = 28-6) per cent. Snout as 
broad as or slightly broader than long, its length 29-0-36-0 (M = 31*6) per cent of 
head ; eye diameter 25-0-31-0 (M = 27-5) ; depth of cheek 22-0-30-6 (M = 26-8) 
per cent. 

Caudal peduncle 13-6-19-0 per cent of standard length, 1-2-1-7 times as long 
as deep. 




Fig. 4. Haplochromis granti, $, B.M. (N.H.) 1956.9. 17.2. Drawn by Miss L. Buswell. 

Mouth usually somewhat oblique ; posterior maxillary tip almost reaching the 
vertical to the anterior orbital margin, or occasionally reaching this line. Lips thick, 
sub-equally developed in a few specimens (e.g. the type), but the upper lip clearly 
thicker than the lower in most. Jaws equal anteriorly, or the lower jaw slightly 
projecting, its length 22-2-30-6 (M = 26-8) per cent of head length and 1-0-1-5 
(mode 1-3) times its width. 

The oblique mouth and unequally thickened lips give an appearance of deformity 
to many specimens. This impression is apparently misleading since there is no 
indication of any impairment to the efficiency of the jaw mechanism, either as a 
mechanical unit or in relation to feeding habits. 

Gill-rakers short, 7-9 on the lower limb of the first arch. 

Scales ctenoid, lateral line with 32 (f.9), 33 (f.n), or 34 (f.9) scales ; cheek with 
3 or 4 (in one specimen 2) series ; 7 or 8 scales between origin of dorsal fin and lateral 
line ; 7 or 8 (rarely 9) between pectoral and pelvic fin bases. 



88 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Fins. Dorsal with 25 (in), 26 (f.17) or 27 (f.i) rays ; anal n (f.7) or 12 (f.21), 
comprising XV-XVII, 9 or 10 and III, 8 or 9 spinous and branched rays for the 
fins respectively. In one specimen the anal fin had been damaged and subsequently 
healed irregularly, giving II, 10 rays. Pectoral shorter than the head, except in two 
specimens where it is of the same length. Pelvic fins extending to the vent in im- 
mature fishes and to the anal fin in adults ; the first ray is proportionately more 
produced in sexually active males. Caudal fin truncate or sub-truncate. 

Lower pharyngeal bone triangular, its dentigerous surface i|— ij times as broad as 
long ; pharyngeal teeth similar to those of H. prodromus. 

Teeth. In the outer series of both jaws, the teeth are similar to those of H. prodromus 
and H. sauvagei ; that is, unicuspid with strongly recurved tips. A few specimens — 
all below 90 mm. S.L. — have some bicuspid teeth situated postero-laterally in both 
jaws. There are 28-46 (mode, ill defined : 36) outer teeth in the upper jaw. 




<S. 



I cm. 

Fig. 5. Mandibular tooth band in H. granti. 



The inner series are composed of tricuspid teeth in most fishes below 90 mm. and 
of unicuspid teeth in larger specimens. An admixture of both types is known from 
three fishes. The teeth are arranged in 2-6 (mode 4) rows in both jaws, but narrow 
to single series laterally. In many specimens the lower tooth band is wider than the 
upper ; antero-medially, the posterior margin of this band is straight or very gently 
curved, thus contrasting with the lower series in H. sauvagei and H. prodromus, 
where the margin is clearly curved (Text-fig. 5). 

Syncranium and associated musculature. The preorbital face of the neurocranium 
is intermediate in form between that of H. sauvagei and the generalized Haplo- 
chromis type. Greatest departure from the condition observed in H. sauvagei and 
H. prodromus is seen in the maxilla, which in H. granti is shorter and more bowed 
in its long axis. Also, the inner face of the posterior limb is markedly concave, 
which results in the outer face appearing more bullate than in other members of 
the " sauvagei " group. The dentary resembles that of H. sauvagei but differs in its 
less rounded, more angular, anterior outline. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES S9 

Shortage of material allowed only two dissections of head musculature to be made. 
The major muscles are distributed as in H. prodromus and H. sauvagei but the origin 
of the adductor mandibulae I is deeper and more fan-shaped in H. granti. In the 
two specimens dissected, the length of this muscle (42 and 43 per cent of head) is 
somewhat greater than in H. prodromus but equal to that in H. sauvagei. 

Coloration in life : Adult males. Ground colour light blue-grey ; branchiostegal 
membrane dusky, especially between the rami of the lower jaw. Dorsal fin blue- 
grey, darkest on the proximal third ; lappets orange-red, as are the spots and streaks 
between the soft rays. Caudal blue-grey, darker on the proximal half ; margin out- 
lined in red ; orange-red spots between the rays. Anal dusky blue-grey, with an 
overall pink flush ; ocelli yellow. Pelvics black, faint pink mesially. Females and 
immature males. Coloration in life unknown. 

Colour in preserved material : Adult males. Ground colour grey or brown ; 
branchiostegal membrane dark grey. An intense black mid-lateral stripe and often 
traces of a lachrymal stripe and 5-7 vertical bars across the flanks. Dorsal, caudal 
and anal fins hyaline or dusky ; pelvics black. Females and immature males. Ground 
colour silver-white, darkest dorsally. An intense mid-lateral stripe and often faint 
indications of an interrupted upper band running between the dorsal fin base and 
the upper lateral line. Seven to nine faint transverse bars are usually present on 
the flanks and caudal peduncle ; no lachrymal stripe. All fins hyaline. 

Distribution. Confined to Lake Victoria. 

Ecology : Habitat. Too few records are available to permit generalization on the 
habitat preferences of H. granti. The twenty-six specimens whose habitat had been 
recorded were caught in littoral zones and in water less than forty feet deep. Most 
localities represented in the collection can be classified either as sandy beaches on 
exposed shores or as exposed coastlines with a hard substrate. The few remaining 
localities are sheltered bays where the bottom is composed of organic mud. 

Food. Twelve of the twenty-six fishes examined contained ingested material in 
the stomach or intestine. In each case only the soft parts of Gastropoda were found, 
except for some Lamellibranchiata shell fragments in one individual. From these 
admittedly few observations it is inferred that H. granti feed in a manner similar 
to that observed for H. prodromus and H. sauvagei. 

Breeding. There is no information on any aspect of the breeding behaviour in 
this species ; all specimens below 90 mm. S.L. were immature. 

Diagnosis. Haplochromis granti differs from other Lake Victoria Haplochromis in 
possessing broad bands of inner teeth (2-6, mode 4, series) in both jaws and by its 
unequally thickened lips, the upper usually thicker than the lower. This latter 
character, together with the oblique mouth and straight posterior margin to the 
inner tooth band of the lower jaw, serves to distinguish H. granti from H. prodromus 
and H. sauvagei. 

Affinities. By virtue of its dentition, H. granti must be included in the H. sauvagei- 
H. prodromus species-group. Other characters probably associated with dentition, 
such as the shape of the premaxilla and dentary, are closely similar in all three 
species. But, despite resemblances in these dental and osteological characters, and 
in the associated musculature, the neurocranial morphology of H. granti has not 



9Q 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



departed so radically from the generalized Haplochromis type. Morphologically 
speaking, the relationship between H. sauvagei and H. prodromus is directly linear, 
whilst that of H. granti is somewhat divergent but with a parallel trophic trend. 



Study material and distribution records 

Museum and Reg. No. 

Uganda 

B.M. (N.H.) 1903.5.30.367 (holotype P. granti) . 
B.M. (N.H.) 1903.5.30.368-369 (paratypes P. 



Locality. 



Bunjako 



Collector. 



Degen. 



B.M. (N.H.) 


1956. 


IO.9. 139 


• 


Bay opposite Kirinya 
Point (Napoleon 
Gulf) 


E.A.F.R.O. 


" 


" 


,, ,, I4O-I4I 


• 


Bugungu (Napoleon 
Gulf) 


n 


» 


" 


,, „ I42-I44 


• 


Beach nr. Nasu Point 
(Buvuma Channel) 


" 


E. African Fisheries Res. Lab. Jinja . 


. 


Pilkington Bay 


1 1 


B.M. (N.H.) 


1956 


.10.9.145-147 . 


. 


Thruston Bay 


it 


,, 


1956. 


9.17.2 • 




Ekunu Bay 


t, 


>> >> 


1956. 


IO.9. I48-I52 




Entebbe, harbour 


it 


>> >» 


,, 


„ „ 153 


. 


Near Busungwe Is. 


a 


" 


" 


„ „ 154 


• 


Busungwe Bay (Kagera 
river mouth) 


. 


t> >» 


,, 


,t ,t 155-157 


. 


Beach near Grant Bay 


a 


" " 


" 


,, ,, 200 


Kenya 


Buka Bay 


" 


11 it 


" 


„ „ 158 


Tanganyikt 


Kisumu 
z 


" 


,, ,, 


,, 


„ „ 159 


. 


Ukerewe Is. 


a 


,, 


,, 


„ „ I6O-I63 




Majita 


,, 



Haplochromis xenognathus sp. nov. 
(Text-figs. 6 and 7) 

The high intra-specific variability of H. xenognathus makes this species of par- 
ticular interest when considering the evolution of monotypic cichlid genera. Some 
of the more aberrant specimens, if studied in isolation, might well be given a status 
equal with the monotypic genera recognized at present. Less extreme individuals, 
on the other hand are not immediately distinguishable from H. sauvagei. 

The modal type tooth-pattern and the usual arrangement of the jaws are, 
however, unlike those of other species in the " sauvagei " group (Text fig. 7). I am 
led to include H. xenognathus in this group because of its " sauvagei "-like tooth 
form and the multiseriate dental pattern. 

The sample provides sufficient intra-specific variation to indicate morphological 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



91 



stages through which the typical specific facies may have passed in its evolution 
from a form similar to the extant H. sauvagei. 

Type specimen. A male, 91 -f- 19 mm. ; from Entebbe harbour. 

Description. Based on thirty-five specimens 80-113 mm. S.L. 

Depth of body 31-2-38-0 (M = 34-8) ; length of head 29-2-35-4 (M = 33-1) per 
cent of standard length. Dorsal head profile usually straight and somewhat steeply 
sloping ; curved in a few specimens. 

Preorbital depth 16-0-20-7 (M = 17-7) per cent head length ; least interorbital 
width 23-5-29-0 (M = 26-8) per cent. Snout from i^-i^ longer than broad, its 
length 31-8-37-8 (M =35.2) per cent of head ; eye 23-2-28-7 (M = 26-0) ; depth of 
cheek 23-2-28-7 (M = 26-0) per cent. 




Fig. 6. Haplochromis xenognathus, J, holotype, B.M. (N.H.) 1956.9. 17.3. 
Drawn by Miss L. Buswell. 



Caudal peduncle 13-8-19-0 (M = 15-9) per cent of standard length ; 1-1-1-7 
(mode 1-4) times as long as deep. 

Mouth horizontal, the posterior maxillary tip reaching, or almost reaching, the 
vertical to the anterior orbital margin ; lips slightly thickened. The lower jar is 
clearly shorter than the upper in 74 per cent of the specimens examined and sub- 
equal to the upper in 26 per cent. Even in this latter group the outermost teeth of 
the lower jaw occlude behind the equivalent upper jaw series. Lower jaw 32-0-38-0 
(M = 34*5) per cent of head, and i-i-i-8 (mode 1-4) times as long as broad. 

Gill rakers short, 7-9 (rarely 10) on the lower limb of the first arch. 

Scales ctenoid ; lateral line with 31 (f.4), 32 (f.14), 33 (f.n), 34 (f.4) or 35 (f.2) 
scales ; cheek with 3 or 4 (rarely 2 or 5) series ; 6 or 7 (less frequently 8) scales 
between origin of dorsal fin and lateral line ; 7 or 8 (rarely 6 or 9) between pectoral 
and pelvic fin bases. 



92 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Fins. Dorsal with 24 (f.13), 25 (f.19), or 26 (f.3) rays ; anal with 11 (f.9), 12 (f.24) 
or 13 (f.i). One specimen has only two spines, giving a total count of 10 rays. The 
spinous and branched ray counts for the fins are XV-XVII, 8-10 and III, 8-10. 
Pectoral fins shorter than the head. Pelvic fins with first ray produced, extending to 
the anterior part of the anal fin in females and more posteriorly in adult males. Caudal 
truncate. 

Lower pharyngeal bone triangular, its dentigerous surface i^-ij times as broad as 
long. The pharyngeal teeth are slender and bicuspid, those of the median series 
not noticeably enlarged. In one specimen, the lower pharyngeal bone is stout, the 
toothed surface slightly longer than broad and the median teeth enlarged and 
molariform ; this fish also shows a somewhat atypical oral dentition, in that the 
teeth are bluntly cuspidate. 

Teeth. Except in the smallest specimen, unicuspid teeth predominate in the outer 
series, but some weakly bicuspid teeth do occur postero-laterally in both jaws of 
large fishes. In the smallest specimen, the entire outermost series is composed of 
bicuspid teeth. 

The outer teeth, like those of H. sauvagei and other species of the group, are 
relatively stout and have strongly recurved tips. In the lower jaw, the anterior 
teeth are implanted at an acute angle, so that their necks lie almost horizontally ; 
but recurvature of the crown is such that the tip points almost vertically upwards 
(Text-fig. 8). There are from 32-52 (mode 44) outer teeth in the upper jaw. 

Teeth forming the inner series are small and either unicuspid or weakly tricuspid. 
Considerable variation exists in the number of inner rows. In general, teeth are 
disposed in a broad crescent which narrows abruptly at a point almost mid-way 
along the premaxillary limb. Thereafter, there is a single inner row. The antero- 
medial depth of this band varies with the number of tooth rows, of which there are 
from 3-9 in both jaws (modes 7 and 5 for the upper and lower jaws respectively). 
In fishes with markedly disparate jaws, the most posterior inner teeth of the dentary 
do not occlude with the upper series. 

The toothed surface of the dentary is often slightly convex, so that when viewed 
laterally several points on the inner band are higher than the crowns of the outer 
teeth (Text-fig. 8). 

Fishes with narrow tooth bands in both jaws have a dental pattern closely resem- 
bling that of H. sauvagei ; the resemblance to this species is enhanced by the sub- 
equal jaws of these specimens. In contradistinction, other H. xenognathus with 
sub-equal jaws have a broad and specifically typical tooth pattern. 

Syncranium and associated musculature. The neurocranium of H. xenognathus is 
identical with that of H. granti. The shape of the premaxilla varies slightly in relation 
to the number of inner rooth rows, but is otherwise comparable with the premaxilla 
of H. sauvagei. Likewise the dentary is similar in the two species, except that the 
dentigerous surface is inclined forwards and downwards in H. xenognathus. Perhaps 
the most characteristic appearance of this bone is imparted by the almost hori- 
zontally implanted anterior teeth. 

A syncranial skeleton prepared from a specimen with H. sauvagei-\ike facies did 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 93 




0*5cm. 

Fig. 7. Premaxillary and mandibular tooth bands in H. xenognathus. 




Fig. 8. 



I cm. 

Lateral view of the anterior part of the dentary of H. xenognathus. 



94 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

not differ, beyond the limits of individual variability, from that of a typical specimen 
and was clearly distinguishable from that of H. sauvagei. 

Head musculature in H. xenognathus is similar to that of H. sauvagei and H. 
prodromus. The adductor mandibulae I is shorter than in the former species, but is 
equal to that of the latter (33*2-39-4 per cent of head). 

Coloration in life : Adult males. Ground colour dark bronze dorsally, shading to 
grey-bronze ventrally : cheek and operculum with a distinct bronze sheen ; chest 
and branchiostegal membrane bluish-grey. Dorsal fin sooty, with red streaks between 
the soft rays ; lappets red. Caudal dark, upper half with red spots, lower half 
flushed with red. Anal clouded ; ocelli yellow. Ventrals sooty, the first ray 
bluish-white. Live coloration of immature males is unknown. Females. Ground 
colour as in males, but the branchiostegal membrane greyish. Dorsal fin with red 
lappets but lacking the red streaks. Caudal dark yellow ventrally, lighter and 
maculate above. Anal and ventral fins dark olive-yellow, the first pelvic ray 
bluish-white. 

Colour in preserved material : Breeding males. Dark grey, the flank with a faint 
coppery sheen ; a distinct lachrymal stripe. In some specimens five transverse bars 
may be discerned on the flanks. Dorsal, caudal and anal fins grey, the upper third 
of the caudal maculate. Pelvic fins black. Non-breeding and immature males. Ground 
colour silver-grey ; 7-9 distinct transverse bars. Fins as above. Females. As for 
non-breeding males, except that the pelvics are colourless. In some specimens there 
is a fairly distinct mid-lateral stripe. 

Distribution. Known only from Lake Victoria. 

Ecology : Habitat. The few and scattered records indicate that H. xenognathus is 
confined to littoral zones where the substrate is hard. Most specimens in the collec- 
tion were obtained from seine nets operated over exposed sandy beaches. 

Food. Seventeen of twenty-three fishes examined contained ingested material in 
the stomach or intestine : in ten, only the soft parts of Gastropoda were found ; in 
three, Insecta (chironomid and ephemerid larvae) ; in two, bottom detritus (sand- 
grains and plant tissue) ; and in three, unidentifiable sludge. 

Although no observations have been made on living fishes, the presence of gas- 
tropod remains without shells suggests that the feeding methods of H. xenognathus 
are like those of H. sauvagei, H. prodromus and H. granti. 

Breeding. No information is available. The smallest sexually mature fish was a 
female 87 mm. S.L. 

Diagnosis. Haplochromis xenognathus may be distinguished from other species of 
the genus by the following characters : outer teeth with strongly recurved tips, 
those of the lower jaw implanted horizontally or almost so ; inner teeth in the upper 
jaw arranged in a broad, crescentic band (3-7, mode 5 series) ; lower jaw usually 
much shorter than the upper and also with broad bands of teeth (3-9, mode 5, inner 
series). Some individuals closely resemble H. sauvagei both in gross morphology 
and in details of dentition, but may be distinguished by the peculiar implantation 
of their anterior lower teeth and by having a slightly narrower and longer snout ; 
in life the coloration of both sexes is diagnostic. 

Affinities. In many respects the species has departed considerably from the basic 



REVISION OF LAKE VICTORIA H APLO.CHROMIS SPECIES 



95 



" sauvagei " type as represented by the nominate species. Yet, it is apparently with 
H. sauvagei that H. xenognathus shows greatest morphological affinity. The resem- 
blance is most clearly seen in the least typical members of H. xenognathus, but is 
obscured in other forms. These latter individuals seem to indicate that if future 
evolution in H. xenognathus is continued along such lines and is coupled with a 
reduction in morphological variation, then the species could acquire a status equi- 
valent to the monotypic genera recognized at present. 



Study material and distribution records 

Museum and Reg. No. 
B.M. (N.H.) 1956. 10.9. 164-167 
„ „ „ 168 



» „ ,/ 169 

,, ,, 170 
1956.9-17-3 (type) 
1956. 10.9. 171-173 

„ „ „ 174 

„ ,, „ 175 

,, ,, ,, 176-177 

„ „ 178-182 



Locality. 

Jinja 

Bay opposite Kirinya 

Point (Napoleon 

Gulf) 
Beach nr. Nasu Point 

(Buvuma Channel) 
Hannington Bay 
Entebbe, harbour 

Entebbe, Airport beach 
Bugonga, Entebbe 

peninsula 
Katebo 
Beach south of Bu- 

sungwe (Kagera river 

mouth) 



Collector. 
E.A.F.R.O. 



183 

184-191 

192-195 



Tanganyika 



Bukoba 

Majita 

Mwanza, Capri Bay 



DISCUSSION 

In an adaptively multi-radiate species-flock, the differentiation of true phyletic 
relationship from parallel trends is difficult, particularly when the flock, like that of 
Lake Victoria, is oligophyletic in origin. It has been noticed, however, that in many 
adaptive sub-groups some species show greater morphological affinity with one 
another than with other members. This I interpret as indicating that both lineal 
descent and parallel evolution have contributed to the origins of the groups. 

The remarkable uniformity of the distinctively shaped outer teeth in all four 
members of the " sauvagei " complex probably indicates a monophyletic origin for 
the group. Their multiseriate dentition, on the other hand, cannot be considered of 
value in indicating phylogeny. Broad tooth-bands have evolved in several other 
and unrelated species, for example, the algal-grazers H. nigricans and H. nuchi- 
squamulatus, and in certain monotypic genera. 

As an ecologically defined group, the " sauvagei " complex exhibits considerable 
variation in species morphology. There are three distinctive forms, represented by 



96 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

H. sauvagei and H. prodromus, H. granti, and H. xenognathus. The two first- 
mentioned species must be considered nearer the generalized type and the two 
latter as showing progressive but independent divergence. 

When discussing the evolution of Macropleurodus bicolor, I drew attention to the 
possible relationship between this species and Haplochromis prodromus (Greenwood, 
1956a). No conclusion can be reached at present, but the prospective adaptational 
significance of a " prodromus " type cranial anatomy in the evolution of M. bicolor 
cannot be disregarded. That both species should have almost identical feeding 
habits would seem to lend additional weight to this argument. 

Morphological and ecological differences between H. prodromus and H. sauvagei 
are of the slightest order. If their lineal relationship can be accepted, one is tempted 
to consider the species in an ancestor-descendant category. In all probability, present- 
day H. sauvagei differ genotypically and even phenotypically from the presumed 
ancestral type, yet, despite these limitations, it is difficult to imagine a species more 
similar to H. prodromus. Several other such Haplochromis species-pairs are known 
from Lake Victoria. Each, except for their temporal coexistence, would fulfil the 
palaeontological requirements for ancester-descendant relationship. Indeed, co- 
existence, even within one habitat, of species which could be lineally derived appears 
to be a major feature of the Lake Victoria Haplochromis species-flock. The Lake's 
geological history provides a possible explanation of this phenomenon (Brooks, 
1950 ; Greenwood, 1951). During the inter-Pluvial periods of the Pleistocene, Lake 
Victoria was probably reduced to a series of small lakes and swamps. Under such 
conditions a species would be isolated into several discontinuous groups. If some 
surviving populations underwent genie reorganization as a result of isolation, or of 
increased selection pressure, it is possible that they might retain their discreteness 
if brought into contact with the parental stocks when the lakes were joined during 
the succeeding Pluvial period. That the derived and parental species were able to 
coexist even when their ecological requirements were similar, seems to indicate 
drastically reduced selection pressure. 

The third member of the " sauvagei " group, H. granti, is unlike either H. sauvagei 
or H. prodromus. However, the differences lie in characters which could be derived 
by heterogonic growth of certain cranial parts from a species less differentiated than 
H. sauvagei, but possessing the group dental characters. 

From an evolutionary viewpoint, H. xenognathus is undoubtedly the most interesting 
species. When the first few specimens came to my notice, I considered them to be 
members of a distinct and apparently monotypic genus. The degree of morphological 
differentiation of this supposedly new genus was at least equal to that of Hoplotilapia 
retrodens Hilg. When more specimens were collected, however, it was obvious that 
the species was extremely variable and should be retained within the genus 
Haplochromis. The less typical specimens differed only slightly from H. sauvagei, 
whilst the typical fishes were clearly distinct from that species. Thus, it seems 
legitimate to look upon H. xenognathus as an example of a stage through which 
species might pass in the evolution of a genus. 

Ecologically, the " sauvagei " group has entered an adaptive zone unique for 
Haplochromis, but occupied by two monotypic genera, Macropleurodus bicolor and 



REVISION OF LAKE VICTORIA H APLOCHROMIS SPECIES 97 

Hoplotilapia retrodens. All other known predominantly mollusc-eating Haplochromis 
species crush their prey by means of hypertrophied pharyngeal bones and dentition. 
If, in conclusion, one considers the morphological and consequent ecological 
adaptations of the H. sauvagei group, the impression is gained of a species-complex 
partially advanced on the path of trophic specialization. From its present peak it 
could supply, and may even have supplied, raw material for further specialization. 

SUMMARY 

1. Haplochromis sauvagei (Pfeffer) 1896, and H. prodromus Trewavas 1935, are 
re-described. 

2. The species H. granti Blgr. 1906, previously synonymized with H. sauvagei, 
is reinstated. 

3. A new species, Haplochromis xenognathus, is described. 

4. Data are given on the ecology of all four species. 

5. Sex-limited polychromatism, involving a piebald female coloration, is described 
for H. sauvagei. 

6. The evolutionary status of the species is discussed. 

REFERENCES 

Brooks, J. L. 1950. Speciation in ancient lakes. Quart. Rev. Biol. 25 : 131. 

Greenwood, P. H. 195 1. Evolution of the African cichlid fishes ; the Haplochromis species- 
flock in Lake Victoria. Nature, London, 167 : 19. 

■ — — 1956a. The monotypic genera of cichlid fishes in Lake Victoria. Bull. Br. Mus. (Nat. 
Hist.) Zool. 3 : 297. 

1 95 6b. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) Part I. 

Ibid. 4 : 225. 

Regan , C. T. 1922. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. 157. 



ACKNOWLEDGMENTS 

I wish to express my gratitude to the Trustees of the British Museum (Natural 
History) for the numerous facilities afforded me in their museum ; to Dr. Ethelwynn 
Trewavas for her constant help and for reading the manuscript of this paper ; and 
to my colleague Dr. Philip S. Corbet who identified the insects recorded from 
stomach contents. 



PLATE 4 

The types of Haplochromis sauvagei (Pfeffer) upper and H. nuchisquamulatus (Hilgendorf) 
lower (see Greenwood, 1956, Bull B.M. (N.H.) Zool. 4, 241), from photographs made in the Berlin 
Museum in 1921. Both specimens may be lost. 



Bull B.M. {N.H.) Zool. 5, 4 



PLATE 4 





PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 
BARTHOLOMEW PRESS, DORKING 



A REVISION OF THE GENERA 
NIDALIA AND BELLONELLA 

WITH AN EMENDATION OF NOMENCLATURE 

AND TAXONOMIC DEFINITIONS FOR 

THE FAMILY NIDALIIDAE 

(OCTOCORALLIA, ALCYONACEA) 



HUZIO UTINOMI 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 5 

LONDON: 1958 



A REVISION OF THE GENERA NIDALIA AND 

BELLONELLA, WITH AN EMENDATION OF 

NOMENCLATURE AND TAXONOMIC 

DEFINITIONS FOR THE FAMILY 

NIDALIIDAE (OCTOCORALLIA, ALCYONACEA) 



BY 



HUZIO UTINOMI, D.Sc. 

(Seto Marine Biological Laboratory, Japan) 



Pp. 99-121 ; 6 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 5 

LONDON: 1958 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY) instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5. No. 5 of the Zoological 
series. 



© Trustees of the British Museum, 1958 




PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued August , 1958 Price Seven Shillings 



A REVISION OF THE GENERA NIDALIA AND 

BELLONELLA, WITH AN EMENDATION OF 

NOMENCLATURE AND TAXONOMIC 

DEFINITIONS FOR THE FAMILY 

NIDALIIDAE (OCTOCORALLIA, ALCYONACEA) 1 

By HUZIO UTINOMI, D.Sc. 



SYNOPSIS 

Redescriptions are given of Nidalia occidentalis Gray, Bellonella granulata Gray and Bellonella 
(Cereopsis) bocagei (Kent) Wright & Studer, based on types in the collections of the British 
Museum. A review of all the known species revealed some diagnostic characters and 
synonymies for the two genera Nidalia and Bellonella, which are here recognized as valid 
separate genera belonging to different families. 

The family Nidaliidae of Gray is re-established for the generally used name Siphonogorgiidae 
and the reasons for this are discussed. The unbranched cylindrical Bellonellids, with somewhat 
contractile calyces, are shown to be essentially primitive, and from them are derived the un- 
branched Nidaliids and more ramose Siphonogorgiids with firm calyces, though not through the 
line of Nephtheidae. 

The concept now outlined of the supposed evolutionary trends in the Alcyonacea is based 
mainly on a consideration of the retractility of polyps as a whole and is contrary to earlier ideas 
of evolution in the group. 

I. INTRODUCTION 

During the course of working on the octocorallian collections in His Majesty's 
Biological Laboratory in Tokyo, and, in particular, when reviewing the Japanese 
species of the so-called genus Nidalia, I have come to recognize a distinctive develop- 
mental tendency between the species, in spiculation and polyp structure, suggesting 
a supposed evolutionary trend from the Alcyoniidae to the Siphonogorgiidae. In 
the meantime, Mr. Frederick M. Bayer of the U.S. National Museum asked me to 
make a comparison between Nidalia occidentalis Gray, type of the genus, and the 
Indo-Pacific forms of " Nidalia ", with a suggestion that the former may be congeneric 
with a Cactogorgia-species from the Indian Ocean, formerly referred to the family 
Siphonogorgiidae. Subsequently to further this work the authorities of the U.S. 
National Museum have very kindly presented a topotypic specimen of Nidalia 
occidentalis preserved in that Museum, together with several specimens of some 
Nephtheid octocorals, to our Laboratory Collection. 

1 Contributions from the Seto Marine Biological Laboratory, No. 307. 

zool. 5, 5. 5 



102 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

The remarkable similarity of the genus Cactogorgia to the older genus Nidalia 
led me to re-examine the types of Nidalia and Bellonella, both of which have been 
considered synonymous by most of the previous authors. Fortunately I have been 
able to do this through the generosity of the British Museum (Natural History) 
and to compare them with the Atlantic and Indo-Pacific (in particular the Japanese) 
forms. In connexion with this, a more critical review of the allied genera formerly 
placed in the families Alcyoniidae and Siphonogorgiidae was necessary. In the 
revision that follows, however, a number of " Siphonogorgia " species were either 
not satisfactorily classified or were left out of consideration, and, in the brief notes 
below, it has been thought desirable to indicate their present status, and where 
possible to add new information. 



II. REDESCRIPTION OF NIDALIA OCCIDENT ALIS GRAY 

(Text-figs. 1-3) 

The following description is mainly based upon a complete specimen presented 
from the U.S. National Museum and partly (spicules only) upon a fragment of the type 
material in the British Museum (Nat. Hist.). 

Material Examined, (i) A specimen labelled as " USNM 50398 ", from 
Pelican St. 169-7 : 28 24-5' N., 8o° 03-0' W., east of Cape Canaveral, Florida, 
45 fms. Jan. 18, 1940. 

(2) Two fragments from the holotype in the British Museum, from off Montserrat, 
West Indies ; depth unknown. 

Description. The colony from east of Cape Canaveral, Florida, is attached to 
a sedentary polychaete tube, on which a young colony of a Muriceid gorgonid, 
Thesea grandiflora, was also living. It is torch-like in form, consisting of a barren 
rigid stalk tapering downwards, and an expanded head-like polyparium covered 
with a number of large verrucae (more than 40) close together at the slightly convex 
summit. The total height is 22 mm., of which about 17 mm. belongs to the stalk 
proper ; the polyparium is 9 mm. in diameter and about 5 mm. high in the middle. 
The largest verruca at the summit of polyparium is about 1-5 mm. in diameter and 
about o*88 mm. in height, and as its tip is blunt, the margin appears rounded. 

The anthocodiae are completely withdrawn into the verrucae. They show a 
well-developed armature at their head. It consists of the eight points and a number 
of collaret rows transversely arranged below, both of which show considerable 
irregularity in the size and arrangement of spicules. In more regular arrangement 
the anthocodial spicules may be counted about 4-5 pairs en chevron in each point 
and up to about 15 transverse rows in the collaret. 

The tentacles, which are simply infolded over the mouth in the contracted condi- 
tion, are probably up to 2 mm. in length when fully exserted and bear about 7-8 
pairs of long pinnules. Their aboral surface is densely packed with small, slightly 
roughened, rodlets which tend to be arranged en chevron and become larger towards 
the base, whereas the pinnules are apparently devoid of spicules or have a few very 
minute rodlets. 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 103 




Fig. 1. Nidalia occidentalis Gray : a, a specimen from East of Cape Canaveral, 
Florida, 45 fms.; b, anthocodial spiculation in two points and in a tentacle ; c, tip of two 
verrucae, with a polyp extended. Note a number of minute scale-like sclerites inside 
the tip of verrucae. 



The middle part of the introvertible neck zone (" the introvert " of Deichmann) 
is wholly bare just below the collarets, while at its base, lining the tip of the verrucae, 
there are numerous very small, colourless, oval scales which are irregularly set, 
but roughly arranged in the eight interseptal tracts. 

The verrucae, i.e., the unretracted thickened part of the polyps, are supported 
by very firm walls densely packed with large stout spindles similar to those of the 
stalk. These spicules are longitudinally disposed close together, and are not grouped 
in eight regions. 

The stalk is deeply furrowed longitudinally on the surface due to the longitudinal 
arrangement of large spicules, which are all strongly tuberculated spindles up to 
2*5 mm. in length, covered with compound tubercles, simply or in clusters. The 
coenenchyme is also densely spiculose, since the canal- walls are very compact with 

zool. 5 , 5 . 5 1 



io 4 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 




mm 



■10.2 



mm 



Fig. 2. Nidalia occidentalis Gray : a and b, part of larger spicules from stalk rind ; 
c, typical spindle of stalk rind ; d, typical spindle of verrucae ; e, anthocodial spiculae ; 
f, flat rodlets from tentacle ; g, scales from introvert ; h, side view of tentacle with dense 
arrangement of spicules (from the type specimen of Gray) . Scale on the left applies to 
a-g, and that on the right to h only. 



similar spindles longitudinally disposed, without any sign of the boundary or the 
difference in size and shape of spicules separating the interior from the surface layer. 
The colour of the specimen is orange-yellow in alcohol, paler towards the base 
of the stalk, due to the degree of preservation of the colour of spicules themselves. 
The anthocodial spicules, together with those of introverts are, however, colourless. 

Measurements of Spicules (in mm.). 

Anthocodial spicules : 
Point — warty spindles with simple warts. 0-36 x 0-035 i °*4 X '°35- 
Collaret — do. 0-4 x 0-05 ; 0-56 x 0-05 ; o-6 x 0-035. 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 105 

Tentacle — flat rodlets with jagged furniture, o-i x 0-036 ; 0-12 X 0-03 ; 0-14 
X 0-028. 

Introvert — scales with scalloped edges. 0-06 x 0-028 ; 0-075 x 0-028 ; 0-09 x 
0-036. 

Stalk spicules : 

Verruca — multituberculate spindles, i-o x 0-23 ; 1-2 x 0-17 ; 1-5 x 0-3. 

Stalk (incl. surface and interior) — do. 1-4 x 0-17 ; 1-5 x 0-29 ; 2-0 x 0-32. 




I 1 1 1 1 1 0.5 



mm 





C^ 



0.2 mm 



Fig. 3. Nidalia occidentalis Gray : a, spindle from stalk rind ; b, anthocodial spindles ; 
c, scales from introvert. All spicules from the type specimen in the British Museum 
(Nat. Hist.). 



Remarks. This specimen as described above corresponds exactly with the type of 
Gray in details of the structure and spiculation ; some spicules of the latter are 
shown in text-fig. 3 for comparison. Deichmann (1936, p. 56) described an additional 
species, Nidalia rigida, but the two are so alike that it is not possible to separate 
them from each other specifically. 

Cactogorgia simpsoni (taken at " Siboga " St. 289) which was fully described and 
figured by Thomson & Dean (1931, p. 184) is, curiously enough, in general agreement 
with this Nidalia occidentalis, in spite of the considerable distance separating their 
localities. 

In particular, the occurrence of numerous small oval scale-like sclerites in the neck 
zone or " introvert ", a character which had not been noted in the remaining species 
of Cactogorgia, proves the former to be identical with the latter. It is indeed wonderful 
that only Mr. Frederick M. Bayer (personal communication) seems to have noticed 
this similarity. 



106 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

Type Locality. Off Montserrat, West Indies. 

Distribution. Atlantic coast of North America and West Indies from off South 
Carolina to off Barbados, in 38-170 fms. (Deichmann, 1936 ; Bayer, 1952 ; 1954a ; 
as Nidalia occidentalis) ; off Timor, Arafura Sea, 112 m. (Thomson & Dean, 1931, 
as Cactogorgia simpsoni). 

III. REDESCRIPTION OF BELLONELLA GRANULATA GRAY 

(Text-fig. 4) 

Bellonella granulata Gray, the type species of Bellonella, was only briefly described 
from Bellona Reef, north-west coast of Australia, with only a figure of the total 
animal (Gray, 1862 ; p. 35). Since then, it has not been found again except for an 
enigmatic record by Thomson & Dean (1931) from " Siboga " St. 240, in the Banda 
Sea, at a depth of 9-45 m., which is only a moderate distance from the type locality. 

As regards the spiculation, Gray mentions only that "It has some characters in 
common with my genus Nidalia but differs from it in the surface of the coral being 
minutely granular, and not spiculose." This very brief account led all later students, 
especially Kukenthal, to misinterpret the status of the genus in the classification 
of this group. Therefore, the re-examination and fuller description of Gray's type 
specimen seemed highly desirable, and this was now made possible through the 
re-examination of material from the British Museum. 

Material Examined. Spicules of type material on two mounted slides. 

Description. The type specimen itself could not be examined by me, but, as 
inferred from the original figure given by Gray (1862, p. 34), the colony consists of 
a cylindrical stalk somewhat expanded at base and a capitate polyparium. A number 
of polyps, crowded together at the top of hemispherical head, apparantly resembling 
those in the preceding Nidalia occidentalis, are completely retractile, and when at 
rest their verrucae appear 8-lobed at the tip as in most of the other Bellonella species, 
and not truncated. 

The outer surface of the stalk without polyps is minutely granular according to 
Gray. This granular appearance depends upon the dense covering of minute spicules, 
some of which are figured here for the first time. The spicules are slender thorny 
spindles, or clubs with more spiny heads derived from spindles. The spindles are 
usually longer than the clubs, and they are all transparent and colourless, though 
some retain still a slightly rosy hue around the axis (see Text-fig. 4B) . 

According to Gray, " the base of the polyps is strengthened with very minute 
spicula, placed in a longitudinal series parallel to each other." These spicules are 
also transparent, slightly rosy to yellowish in colour and somewhat club-like in 
form, bluntly headed (see Text-fig. 4A). 

Measurement of Spicules (in mm.). 
Stalk spicules : 

Spindles — 0-17 x 0-04 ; 0-25 x 0-07 ; 0-28 x 0-05 ; 0-32 x 0-035. 

Clubs — o-i x 0-05 ; 0-14 x 0-05 ; 0-17 x 0-05 ; 0-25 x 0-05. 
Polyp spicules : 

Clubs — 0-16 x 0-035 i 0#I 9 X 0-05 ; 0-25 x 0-05 ; 0-28 x 0-05. 

Remarks. According to Thomson & Dean (1931), a small specimen, taken at 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 107 

" Siboga "St. 240, which was recorded under the name " Nidalia granulata (Gray) " 
is about 1 -5 cm. in height, with a maximum diameter of 5 mm. and ochraceous in 
colour. The coenenchymal spicules are said to be " minute double spheres, knobbed 
capstans, a few warty rods " and practically no spindles are included. It is thus 
beyond doubt that they have erroneously identified this specimen, without making 
reference to the type specimen. Their description without figures is too inadequate 
to permit any other suggestions. 





0.1 mm 



Fig. 4. Bellonella granulata Gray : a, spicules from the polyps ; b, spicules from the 
stalk. All spicules from the type specimen in the British Museum (Nat. Hist.). 

Distribution. Type locality only — Bellona Reef, north-western coast of Australia, 
17 fms. 



IV. REDESCRIPTION OF BELLONELLA BOCAGEI WRIGHT & STUDER 

(Text-figs. 5-6) 

The following description is based on fragments of the material of Bellonella bocagei 
(Kent), collected by the Challenger from the west of Azores, and now in the collections 
of the British Museum. 



108 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

Description. The specimen is excellently figured by Wright & Studer (1889, p. 
241 ; pi. 37, fig. 2) as here reproduced in text-fig. 5A, but their description is not 
sufficiently detailed for recognition of the species. 

The specimen figured forms a cylindrical colony arising from a flat extended base. 
As measured from the original figure, it rises from the base to a height of about 
6 cm., with a diameter of about 8 mm. in the middle of the stalk. The upper half 
is rather loosely covered with large cylindrical polyps, and the lower half is apparently 
bare, bearing no polyps at all. 







B 




H 0.5 mm 



Fig. 5. Bellonella bocagei (Kent) : a, a Challenger specimen (redrawn from Wright & 
Studer, 1889) ; b, anthocodia with partially contracted anthostele. 



When extended, the polyps may attain a length of about 2 mm., with a diameter 
of about o-8 mm. at the head. At the base each forms an 8-lobed, low verruca which 
is thick-walled and spiculiferous. The tentacles bear slender, curved or spiny, 
flat rodlets on the aboral side. The anthocodial armature consists of 8 double rows 
of steeply-converging slender spindles with low warts and bluntly ended. Below these 
the similar spindles are arranged in about 10 transverse rows, and they become 
sparser and smaller in size in the eight interseptal tracts of the neck zone down to 
the basal calyx (Text-fig. 5B). 

The stalk cortex is closely packed with long, spiny spindles or shorter, spiny 
clubs thickened at upper end. The coenenchyme contains more slender spindles 
with high warts. 

All these spicules are usually transparent and colourless, but some reddish ones 
are found on the cortex of polyparium. 

Measurements of Spicules (in mm.). 

Anthocodia — 0-25 X 0-05 ; 0-35 x 0-05 (Text-fig. 6a). 

Tentacle — 0-09 <-^o-i4 x o-oi ~o-02 (Text-fig. 6b). 

Neck zone — 0-035 ~ 0*055 X o-oi '^0-03 (Text-fig. 6c). 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 109 




Fig. 6. Bellonella bocagei (Kent) : a, anthocodial point spicules ; b, tentacle spicules ; 
c, spicules from neck zone ; d, spicules from stalk rind ; e, coenenchymal spicules in 
canal- walls, All are drawn from a specimen in the British Museum (Nat. Hist.). 



Stalk cortex — 0-09 x 0-035 '> 0-12 x 0-05 ; 0-23 x 0-035 (Text-fig. 6d). 

Coenenchyme — 0-09 ^0-25 X 0-009 ^ 0-017 (Text-fig. 6e). 

Remarks. This species was originally described by W. Saville Kent (1870, 
p. 398) under the name Cereopsis Bocagei gen. nov. et sp. nov., from specimens 
taken off Setubal, Portugal in 15 fms. Wright & Studer (1889) identified this specimen 
with Kent's species and transferred it to Gray's genus Bellonella, together with 
Nidalia atlantica Studer (1878, p. 635) and Itephitrus 1 speciosus W. Koch (1886) 
recorded from the neighbouring waters as synonyms. This procedure was followed 
by Putter (1900) who reviewed the hitherto known species of Bellonella. Most of 
the later authors, however, regarded this species as a member of Gersemia (Kukenthal, 

1 Erroneously called Iphethyrus or Iphythyrus by Wright & Studer and May respectively. 



no A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

1906a, b, 1907 ; Thomson, 1927) or as a member of Alcyonium (Molander, 1915 ; 
Deichmann, 1936). 

Apart from a problem concerning the distinction between the families Alcyoniidae 
and Nephtheidae from which such different opinions were probably derived, I 
came to the conclusion, by a close comparison of all Japanese species of Bellonella 
with other related genera of both families, that its assignment to the genus Bellonella 
is highly advisable as proposed by Wright & Studer. 

Distribution. Off Setubal, Portugal, 15 fms. (Type locality) ; off Senegal, 
tropical West Africa, 115 fms. (" Gazelle " St.) ; west of the Azores, 450 fms. Chal- 
lenger St. ; Rolas, Gulf of Guinea ; Azores, 845 m. 1 (" Prince Albert I " St. 584). 



V. VALIDITY OF THE GENUS BELLONELLA AND ITS SYNONYMY 

As mentioned above, the genus Bellonella was created by Gray (1862) for a single 
species B. granulata, and later (1869) placed in his family Bellonelladae near the 
Xeniidae. At that time he recognized an obvious difference in spiculation separating 
it from his earlier genus Nidalia although he did not go into details. 

His systematization of various octocorallian genera has not, in general, been 
accepted in modern systems of classification, and his views on the distinguishing 
characters between Bellonella and Nidalia have been opposed by most later authors. 
Wright & Studer (1889) a * ^ TS ^ suggested the probable identity of both the genera, 
though actually treating them separately. In recent years the two genera have been 
united by May (1900), and in particular, by Kiikenthal (1906a, b), who was convinced 
that some species contained in the genus Bellonella (Nidalia in his sense) should be 
retained in the Alcyoniidae, while others like Gersemia and Capnella should be 
transferred to the Nephtheidae. 

Among recent authors, only J. S. Thomson (1910, 1921) expressed a doubt as to 
whether Gray's Nidalia and Bellonella are really identical, and referred two South 
African species to the latter genus as B. studeri n. sp. and B. rubra Brundin. In 
addition to these, he recognized Pfeffer's Metalcyonium as a distinct genus from Bel- 
lonella, referring three species to it. 

Prior to him, Kiikenthal (1906a) divided the genus Alcyonium Linnaeus into 
three subgenera, Alcyonium s. str., Metalcyonium and Erythropodium (later changed 
as Par erythropodium). Although this attempt has not been followed by most other 
authors, especially Molander (1915), it is undoubted that Metalcyonium is a unique 
group embracing the species which are clavate, capitate or mushroom-shaped and 
ordinarily unbranched in form. Most of them were known from the subantarctic 
region (Patagonia and South Africa), but later a few were recorded from Amboina, 
East Indies (Burchardt, 1902) and northern Japan (Yamada, 1950). 

Judging from the descriptions given by Pfeffer (1889) and Kiikenthal (1906a), 
and also from photographs published by Molander (1929, pi. IV, fig. 9), Metalcyonium 
clavatum Pfeffer, which is the type of the genus, seems to be very different from 
others since referred to the genus (or subgenus) in its form of growth. Although I 

Thomson's identification is still in doubt (see Deichmann, 1936 ; p. 51). 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA in 

have no personal knowledge of this species, its close affinity with the genus Bellonella 
cannot be denied. 

Kukenthal (1906a, b), in dividing the family into two subfamilies, Nidalliinae 
(sensu Kukenthal) and Alcyoniinae, emphasized that the canal system is direct and 
partly indirect in the former, while indirect in the latter. Such a difference was, 
however, strongly rejected by Molander (19 15). 

According to Molander, Metalcyonium clavatum often shows a sign of slight 
division of the polyparium into side branches (or lobes) . Here I only wish to point 
out that on investigating the Japanese species of Bellonella such examples as M. 
clavatum could be observed normally or abnormally (Utinomi, 1957). 

Even if this M. clavatum can be regarded as belonging to Bellonella on account 
of the unbranched cylindrical form of growth, this cannot be applied to other 
mushroom-like forms such as M. capitatum Pfeffer, patagonicum May, molle Burchardt 
and novare Kukenthal, as well as other unbranched alcyoniids tentatively referred 
to the genus Alcyonium in the widest sense (for example, see J. S. Thomson, 1910 ; 
Yamada, 1950 and Tixier-Duri vault, 1954). 

If this re-grouping is actually justified as limited by the type designation for 
Pfeffer's genus Metalcyonium, these capitate forms are left without a genus and there- 
fore require a new genus or subgenus name. But for the present, the differences 
between these unbranched, either cylindrical or capitate, forms of Metalcyonium 
and many of lobate or branched forms of the true Alcyonium s. str. are so vague, 
that only by a complete revision can their status be decided. 

In this revision below, though admittedly not complete, the position of the species 
which have been referred to Nidalia (Bellonella) or other genera is considered chrono- 
logically and the conclusion reached earlier for others are re-stated briefly. 

Bellonella granulata Gray (1862) is designated by monotypy as the type of a valid 
genus Bellonella Gray, 1862. 

Cereopsis Bocagei Kent (1870) is, as mentioned above, referable to Bellonella, 
following Wright & Studer (1889) and May (1900). Its assignment to either Alcyonium 
or a Nephtheid genus Gersemia, as proposed by later authors, is not adequate. 

Another species C. studeri, described by von Koch (1891) from Naples, Italy, 
was referred to Nidalia by May, and then to Gersemia by Kukenthal. This species 
was later re-discovered and fully described by Thomson & Dean (1931) from the 
East Indies and by Stiasny (1941) from Naples, under the original name. Very 
recently I had a similar specimen, referable to this species, from Sagami Bay, 
Japan (unpublished) and noted some remarkable characters generically distinguish- 
able from the type species of Cereopsis. Therefore, the genus name Cereopsis cannot 
be used for both bocagei and studeri as a synonym of Bellonella, though it was later 
replaced by the substitute name Cereopsida Strand (1928). Koch's studeri appears 
to be generically distinct from the type species bocagei, and therefore requires 
a new genus, for which the new name Kochella is here proposed. Detailed discussion 
as to this form will appear in another paper. 

Nidalia atlantica Studer (1878, p. 635) 

Itephitrus speciosus W. Koch (1886, p. 1) 
Both species are probably synonymous with Bellonella bocagei (Kent), together with 
the Challenger specimen described above. 



H2 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

Nidalia arctica Danielssen (1887, p. 119) 
This species, together with Organidus nordenskjoldi and Kryst alio fanes polaris succeed- 
ingly described in the same paper, are probably not Bellonellids but may be merely 
young or stunted specimens of Gersemia fruticosa (Sars) (= G. rubiformis, sensu 
Madsen, 1944). 

Bellonella variabilis Studer (1901, p. 25) 

(= Rhodophytum variabile Studer, 1890, p. 89) 
Kiikenthal (1906a, b) referred this species to Gersemia, but Molander (1915) who 
studied Studer's original type, and Deichmann (1936) consider it a stunted specimen 
of Alcyonium glomeratum Hassall with the least development of polypiferous lobes. 

Bellonella rubra Brundin (1896, p. 6) 

Bellonella cinerea Brundin (1896, p. 8) 
Both species first recorded from Japanese waters distinctly belong to Bellonella. 
The identity of the latter with the former, as proposed by Kiikenthal, is still open 
to question and must await the discovery of more material. 

Bellonella rigida Putter (1900, p. 448) 

Eleutherobia japonica Putter (1900, p. 449) 
The latter is undoubtedly a synonym of the former. This is a sand-dwelling form, 
where the stalk is often rounded at the base, apparently as in the pennatulids, such 
as Cavernularia and Veretillum. In fact, Thomson & Rennet (1927, p. 143) carelessly 
included it in the report on the Japanese species of pennatulids. 

Nidalia foliacea May (1900, p. 101) 
Probably identical with a Nephtheid Capnella imbricata (Quoy & Gaimard). 

Bellonella indica Thomson & Henderson (1905, p. 274) 
Probably a valid species with coenenchymal spicules of capstan type. 

Bellonella studeri J. S. Thomson (1910, p. 550) 

Bellonella rubra Brundin (J. S. Thomson, 1910, p. 554) 

Metalcyonium clavatum Pfeffer (J. S. Thomson, 1910, p. 556) 
These three species recorded from South Africa are probably valid species of Bel- 
lonella. But the second species may be different from the species occurring in Japan. 
According to Molander (1929), the third is not the same as the typical species from 
South Georgia. 

Nidalia rubra (Brundin) (Tixier-Duri vault, 1954 ; p. 127) 

Nidalia morifera Tixier-Duri vault (1954, p. 128) 
The former, though briefly described, may be similar to the species of the same 
name from South Africa, which is mentioned above. The latter is a peculiar Bel- 
lonella, closely resembling B. grayi (Thomson & Dean, 193 1) in having an indistinct 
sterile stalk and in having no spicules in the anthocodiae. 

Apart from these, a number of new species have been described by Kiikenthal 
(1906&), Nutting (1912) and Thomson & Dean (1931), all referring to Nidalia. The 
majority of them should be placed in the genus Bellonella as valid species, but only 
a direct comparison with their types will decide it. Further information as to the 
synonymies and affinities is given in my recent paper reviewing Japanese species of 
Bellonella (Utinomi, 1957). 

Consequent upon re-examination of the types of both Nidalia and Bellonella and 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 113 

a review of all previous records referred to both genera and related genera, a revised 
diagnosis of the genus Bellonella Gray is given below : 

Diagnosis. Alcyoniids whose colonies are cylindrical or subcylindrical. Colony 
with a stalk and an unbranched (scarcely slightly-lobed) cylindrical polyparium. 
Polyps large, monomorphic, fully-retractile within 8-lobed, or truncated, calyces. 
Gastric cavities of all polyps closely fascicled, extending to base. Anthocodiae, with 
or without, 8-chevroned rows of spicules. Coenenchymal spiculation sparse but dense 
in outer cortical layer. Spicules : spindles, rods, clubs and capstans. Usually 
vividly coloured. Living in deep waters of all the oceans. 

Type species : Bellonella granulata Gray (1862). 

VI, SYSTEMATIC POSITION OF THE GENUS NIDALIA AND 
ITS RELATIONSHIP WITH SIPHONOGORGIIDS 

The genus Nidalia was erected by Gray (1835) to contain a single species, N. 
occidentalism from off Montserrat, West Indies. The original description given by 
Gray is quite insufficient for the diagnosis of the genus and species, and there was 
no figure or description of the spicules. Accordingly, regrettably enough, Studer 
(1901) and Kukenthal (1906a, b) merged the genera Nidalia and Bellonella as synony- 
mous. The former authority used the name Bellonella, while the latter on the contrary 
the older name Nidalia as the generic name. Nevertheless, Kukenthal actually 
neglected to consider the name of Nidalia occidentalis , which is the type species, 
in recording all of the hitherto known and undescribed species of the genus {Nidalia 
or Bellonella). It is possible that he might have denied its actual existence. This 
confusion might be due to the lack of exact knowledge of both the types, and the 
superficial resemblances of the colonies, based mainly on the imperfect descriptions 
of Gray. 

Since the original descriptions, both the type species were not found again for 
many years. Deichmann (1936) was apparently the first to recognize Nidalia 
occidentalis as a distinct species and genus. At the same time she described another 
species (N. rigida), but her species cannot be regarded as distinct and she did not 
put forward any further comparison with the Indo-Pacific forms of " Nidalia ". 
Accordingly, more recent workers, including myself, have followed Kukenthal 
in considering Bellonella a synonym of Nidalia. 

As has already been remarked in the preceding chapters, Nidalia occidentalis 
Gray differs considerably from Bellonella granulata Gray and the allied forms. The 
differences justify separating the two as different genera and even different families, 
the former as a member of " Siphonogorgiidae ", and the latter to the Alcyoniidae. 

The diagnostic salient characteristics of Nidalia occidentalis may be summarized 
below : 

The colony is torch-like (instead of cylindrical in a strict sense), unbranched, 
with an expanded hemispherical capitulum covered with large crater-like calyces. 
Calyces are not 8-lobed, but are truncated at the tip, and thick-walled, being closely 
packed with large, multituberculate spindles arranged longitudinally. Coenenchymal 
spiculation in the stalk is more rigid, giving the whole colony a brittle consistency. 

These characters indicate its closest affinity with the " Siphonogorgiidae " amongst 
families of the Alcyonacea. 



ii 4 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

In searching in literature I have not seen it mentioned in any paper, where it might 
be expected to be found. As mentioned above, only Mr. Frederick M. Bayer (personal 
communication) seems to have been aware of a remarkable similarity between 
Nidalia occidentalis and Cactogorgia simpsoni. 

The genus Cactogorgia was established by Simpson (1907 ; see also, Thomson & 
Simpson 1909, pp. 143-150) for three species, showing different growth forms, 
collected by the R. I. M. S. ship Investigator in the Indian Ocean. They are C. 
celosioides from Andamans (depth unknown), C. expansa from off Cape Comorin in 
38 fms. and C. alciformis from Andamans and off Arakan coast from 13 fms. Since 
then, three more species have been recorded, also from the Indian Ocean. They are 
C. lampas Thomson & Mackinnon (1910) from the Seychelles from 37 fms., C. agarici- 
formis Simpson (1910) from an unknown locality and C. simpsoni Thomson & Dean 
1931) from " Siboga " St. 289 (9 0-3' S., 126 24-5' E., 112 m., SE. of Timor). 

Notwithstanding the differences in the shape of the colony and the anthocodial 
armature, all of these species agree well with one another in not having any definite 
branching and in having a marked distinction into sterile trunk and polyp-bearing 
portion, and in showing dense spiculation of the canal- walls, as well as in the rind. 
In addition, all spicules are very large, stout, highly-tuberculate spindles of the 
Siphonogorgia-type, and the tentacle spicules are minute, scale-like ; the anthocodiae 
show the definite " crown and points " armature arrangement and are completely 
retractile within the thick- walled verrucae, as in Nidalia occidentalis and many 
of Siphonogorgia species. They are generally yellow to orange coloured. Therefore 
I do not hesitate to regard the two genera as congeneric, and in doing so note that 
Nidalia has priority over Cactogorgia as the generic name. 

As regards the systematic position of this Cactogorgia and also Agaricoides (Simp- 
son, 1905 ; Thomson & Henderson, 1906), opinions differ between the British and 
German authorities. Originally Simpson placed both among the Siphonogorgiinae 
(as a subfamily of the Nephtheidae) on account of the rigid consistency of the colony 
caused by the dense spiculation in the coenenchyme. Among recent authors, for 
example, Chalmers (1929), Hickson (1930) and Bayer (1956) consider that they are 
more closely related to the family Siphonogorgiidae than to any other. 

Kiikenthal (1896) raised the subfamily Siphonogorgiinae to the rank of a separate 
family as Siphonogorgiidae in the belief that Siphonogorgia is intermediate in form 
between the Nephthyidae (= Nephtheidae) and the Gorgoniidae (= Gorgonacea, in 
particular, Scleraxonia) . Nevertheless, later (19066, 1910) he pointed out the 
dissimilarity of Simpson's two new genera to the gorgonids, suggesting only a near 
relationship of Agaricoides to Nidalia (= Bellonella) macrospina Kiik. and that of 
Cactogorgia to Nidaliopsis pygmaea Kiik., both belonging to the Alcyoniidae. 

From an examination of N. macrospina from Japan Kukenthal's suggestion is 
amply confirmed. In this connexion, a note by Thomson & Simpson (1909, p. 135) 
recalls that Siphonogorgia annectens n. sp. " bears a strong resemblance to Nidalia 
macrospina Kiikenthal.' ' This siphonogorgiid is, according to Macfadyen (1936), 
probably synonymous with Nephthyigorgia pinnata which genus was created by 
Kiikenthal (1910) for three Australian species showing a poor internal spiculation 
in the stem. This view also may possibly be right. 

If these closer relationships to either of the two families Siphonogorgiidae and 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 115 

Alcyoniidae (instead of Nephtheidae as generally accepted) are actually justified, 
then we are faced with the alternatives of expanding the definition of either of the 
two families generally accepted or erecting a new separate family for these unbranched 
forms. 

A search of the literature on Siphonogorgia Kolliker (including Chironephthya 
Wright & Studer), comprising a large number of species, reveals that the hardness of 
coenenchymes due to the dense internal spiculation, given by Wright & Studer 
(1889) as diagnostic, is quite untenable. As can be deduced from a list of hitherto 
known species of Siphonogorgia given by Thomson & Dean (1931, pp. 153-166), 
the coenenchymal spiculation especially in the canal- walls is very variable, and in 
another closely related genus, Nephthyigorgia, the distribution of spicules is largely 
confined to the outer layer (usually called cortex or rind) . 

The rigid consistency and brittleness of the colony which characterize Siphono- 
gorgia and allied genera are mainly due to the thick wall formed by a definite close 
arrangement of longitudinally-disposed, large spindles, regardless of internal spicu- 
lation whatever. Therefore, the colonies are decidedly different structurally from those 
of the Gorgonacea, since no spicular axis as seen in the Scleraxonian gorgonids is 
formed in the interior, although there may be a superficial resemblance in the mode 
of growth. 

In both the genera mentioned above, Nidalia (Cactogorgia) and Agaricoides, 
the colonies are generally unbranched, like alcyoniids such as Bellonella and Antho- 
mastus, but the spiculation of the cortex and the structure of polyps are more closely 
related to the Siphonogorgiidae than to the Alcyoniidae. Thus it seems better to 
include both under the former group as a special subfamily rather than to erect a 
separate family. 

In doing so, however, the generally used family name Siphonogorgiidae (pro 
Siphonogorgiaceae Kolliker, 1875, p. 22) should be displaced by the name Nidaliidae 
(pro Nidalidae Gray, 1869, p. 127), since the latter has priority over the former as 
the family name. Gray's diagnosis, that is " Coral simple or branched ; stem cylindri- 
cal, cartilaginous, with a crustaceous skin and imbedded spicules. Polypes on the 
upper surface of a hemispherical head, with prominent large conical polype-cells ; 
stem and polype-cells covered with fusiform spicules " is applicable to this family 
without much alteration of diagnostic characters. 

Below I propose a new system of classification based on the revised examination 
of the type specimen of Nidalia and reconsideration of the diagnostic characters of 
the allied genera, formerly assigned to the Siphonogorgiidae, in recognition of the 
distinctive evolutionary trends they display. 

Family Nidaliidae Gray, 1869 

Nidalidae Gray, 1869, p. 127. 

Siphonogorgiaceae Kolliker, 1875, p. 22 ; Klunzinger, 1877, p. 48 (as subfamily of 

Alcyonidae in Alcyonacea). 
Siphonogorginae Wright & Studer, 1889, p. 226 ; Thomson & Henderson, 1906, 

p. 11 ; Thomson & Simpson, 1909, p. 124 (as subfamily of Nephthyidae) . 
Siphonogorgiidae Kiikenthal, 1896, p. 133 ; May, 1900, p. 171. 
Siphonogorginae (sic) Hickson, 1903, p. 487 (ranked however as a family). 



n6 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

Siphonogorgiidae Kukenthal, 1906&, p. 68 ; Harrison, 1909, p. 31 ; Hickson, 1930, 

p. 244 ; Thomson & Dean, 1931, p. 149 (part) ; Bayer, 1956, p. 188 (part). 
Not Nidaliinae Kukenthal, 1906a, p. 29 (as subfamily of Alcyoniidae). 
Not Nidalinae (sic) Kukenthal, 19066, p. 19 (as subfamily of Alcyonidae). 

Emended Diagnosis. Colonies unbranched or tree-like, branched with stiff, 
cylindrical branches. Outer surface very roughened, closely-packed with large, 
multi-tuberculate spicules longitudinally-disposed, which give the colony a rigid 
brittle consistency. Anthocodiae completely or partially retractile, within densely 
spiculose tubulo-conic verrucae, more or less projecting above the surface. 
Anthocodial armature with symmetrical " points and crown " arrangement. 

Subfamily Nidaliinae nov. 

Unbranched, or slightly branched only at base ; colonies with a marked distinction 
into trunk and polyp-bearing portion ; densely spiculose throughout ; polyps 
completely retractile. 

For the present, only two genera Nidalia Gray (= Cactogorgia Simpson) and 
Agaricoides Simpson are known as belonging to this subfamily. 

Subfamily Siphonogorgiinae Kolliker 

Profusely or feebly branched, tree-like colonies not distinctly separable into trunk 
and polyp-bearing branches ; canal-walls filled, densely or loosely, with spicules 
similar to those of rind ; polyps singly or closely-arranged on stem and completely 
or partially retractile. 

Siphonogorgia Kolliker is a representative genus among this group. N ephthyigorgia 
Kukenthal is also probably distinct, but Dactylonephthya Thomson & Simpson seems 
unique in the absence of distinction between the anthocodiae and verrucae, if this 
proves to be correct. Here it is tentatively placed near the position of Siphonogorgia, 
but its re-discovery is much needed. 

The remaining genera, such as Scleronephthya Wright & Studer and Stereacanthia 
Thomson & Henderson, if distinct as genera, may be grouped within the Nephtheidae, 
together with Capnella (= Paranephthya) and Lemnalia. 

VII. EVOLUTIONARY INTERRELATIONSHIP BETWEEN 

ALCYONIIDAE AND NIDALIIDAE 

(SIPHONOGORGIIDAE AUCT.) 

Our knowledge of the interrelationships of various groups and their evolutionary 
trends in the Alcyonaria is very incomplete. Kukenthal (1906a) has discussed the 
evolution of the Alcyonacea, concluding that the Siphonogorgiidae are the most 
evolved group derived from a supposed solitary Haimeia-like ancestor along a line 
Cornulariidae-Xeniidae-Alcyoniidae-Nephtheidae. But this monophyletic view has 
not been supported by later workers (in particular, Molander, 1915). Within the 
Alcyoniidae, Molander supposed two different lines of evolution as regards polyp 
dimorphism (one includes the monomorphic forms, and the other, the dimorphic 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 



117 



forms) and thought that Nidalia (sensu Kiikenthal) was the most primitive genus. 
His view seems to me adequate, but he did not extend his views to mention any 
further evolutionary lines along which they may have evolved. 

As discussed above, Bellonella (namely, Nidalia sensu Kiikenthal) is more primitive 
than the lobated or ramified forms of Alcyonium for reasons already given, and it is 
among this genus that we have to look for forms resembling the ancestral retractiel 
stoloniferans. Nidalia (sensu Gray), on the contrary, is a more advanced or specialized 
form to be placed near Siphonogorgia, although it is ordinarily unbranched in its 
mode of growth. 

The two genera have been treated as synonymous for many years, perhaps because 
of the lack of true knowledge of the original types of the two, or of confusion with 
regard to the species status due to the superficial resemblance of the colonies. 

The most primitive forms, such as B. rubra and B. grandiflora, retain the flaccid 
consistency of the coenenchyme, of which canal- walls are thick but not so densely 
spicular as in the rind. The coenenchymal spicules are of uniform shape and small in 
size, and the anthocodial armature retains many ancestral features seen in stoloni- 
ferans. The retractile polyp is very large, supported by a long stalk with flexible 
armature of small spicules continued downwards as 8 interseptal areas, so that, in 
contraction the short stiffened lower part of the polyp itself, called anthostele, is 
infolded to cover over the retracted anthocodia, forming a 8-lobed wart-like operculum 
by which the mouth is completely closed. 

In these species showing the 8-lobed anthostele (" calyx "), the arrangement 
of spicules on the surface is ordinarily continuous between the anthocodia and 
anthostele, although sometimes interrupted by reducing spicules in the neck zone, 
and the anthostele is more or less contractile, and thus functional as an operculum 
of the polyp itself. 

In the more evolved species such as B. sibogae (= B. macrospina Thomson & 
Dean) and B. macrospina (= Nidalia macrospina Kiikenthal), the cortical spiculation 
is more massive and rigid, with greater stoutness of spicules. The polyps (more 
properly the anthocodiae) , on the other hand, shorten the stalk, reduce the spicular 
armature in the introversible neck zone, and become heavier than the anthocodial 
armature. 

Consequently, the discontinuity between the anthocodia and the anthostelar 
region in spiculation may become more pronounced and a thick-walled, non-contractile 
calyx as a tubular upgrowth from the cortical coenenchyme (or rind) results. The 
calycinal spiculation is not arranged interseptally and the apical orifice cannot be 
completely closed. Here the anthocodia itself, protected by spicular armature, takes 
a part as an operculum, as is usual in most siphonogorgiids and gorgonaceans. 
For further details on the evolutionary trend within the genus Bellonella, see Utinomi 

(1957). 

In the genus Siphonogorgia the arrangement of anthocodial armature is variable 
between species, as pointed out by Chalmers (1929) in discussing the evolution of 
species within the genus. It is also probable that this arrangement is intimately 
concerned with the retractility of the anthocodiae. 

As regards the origin of Siphonogorgia, in establishing the genus, Kolliker (1875) 
regarded it as intermediate between the Alcyonacea and the Gorgonacea in modern 



u8 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

systems of classification. Since then, following him, most later authors generally 
consider that Siphonogorgia and allies are derived from the Nephtheidae or at least 
more related to it than to any other families, a conclusion from which I must most 
emphatically dissent. The generally accepted ideas, in particular Kukenthal's, 
that may have been derived from considering its Nephtheid-like mode of growth, 
associated with the heavier coenenchymal spiculation, and its warm- water distribution 
restricted to the Indo-Pacific, also appear to be untenable. 

The growth form of the colony is, I believe, of limited significance in assessing 
whether a particular form is primitive or advanced, but reflects general trends in 
the group. This may reflect the effects of the external environment, especially of 
water movements, sea temperature and kind of substratum, although there are certain 
basic characteristics which are apparently never modified by the environment. Thus, 
such a slight sign of branching in Nidalia occidentalis as reported by Gray, as well as 
a few indistinctly-lobed examples in some species of Cactogorgia, suggests a probable 
evolutionary trend towards ramified siphonogorgiids. 

In most of the genera within the family Nephtheidae, the polyps, distributed 
scatteredly or in groups on branches, are ordinarily not retractile, forming prominent 
calyces without clear division between the anthocodia and anthostele, and conse- 
quently calycinal spicules are completely continuous with those of the cortical 
coenenchyme of branches and stems. Each polyp is either cylindrical or clavate in 
shape. In the latter the spicules in the anthocodial region are somewhat bilaterally 
symmetrical in arrangement, instead of being radial as in most alcyoniids. 

The retractile polyps in Gersemia and Paralemnalia, as in a few exceptional cases, 
are in all probability a secondary outcome due to the weak development of spicula- 
tion in the neck zone. Similar trends as discussed above may probably be observed 
also within the Gorgonacea. 

As regards another aberrant family called Viguieriotidae Bayer (19546), better 
known as Fasciculariidae Viguier, I have no personal knowledge at present. But it 
is likely that this family may be a complex of allied forms, having in common only 
a densely spiculose stalk (" cup " or " involucre ") into which retract numerous 
polyp-bearing lobes or branches. In other respects they are not related to one another, 
since one Viguieriotes (= Fascicularia Viguier) (the simplest) is closely related to the 
Clavulariidae, one (Paralcyonium) to Bellonella or Nidalia, whereas the remaining 
one (Studeriotes) is more specialized with Nephthea-like ramified twigs and polyp 
armature. Even if these genera may seem to have originated in different ways, there 
are insufficient reasons for refusing to combine them into a special family. 

In conclusion, it need only be mentioned that the retractibility of polyps is indeed 
one of the important characteristics originally bestowed on the Octocorallia and even 
all anthozoans. It is a pity that most octocorallian specialists did not lay enough 
stress on this peculiar character in recognition of various groups within the Octo- 
corallia and their evolutionary trends. 

VIII. ACKNOWLEDGMENTS 

I am very much indebted to the Trustees and Dr. W. J. Rees of the British Museum 
(Natural History) for the gift of some fragments from the Gray's type specimens 
and some alcyonarians in the Challenger Collections mentioned in the text and for 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 119 

much help in publishing this paper. I am particularly indebted to Mr. Frederick 
M. Bayer of the U.S. National Museum, Washington, for his helpful suggestion and 
criticism and also for many duplicated specimens sent to me for comparison. I 
wish to extend my sincerest thanks to Dr. Hirotaro Hattori for the privilege of study- 
ing His Majesty's vast collections to which I owe the opportunity to begin the study 
of the group. 



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1954a. Anthozoa : Alcyonaria. In, Gulf of Mexico, its origin, waters and marine life. 

Fish. Bull., U.S. 89 : 279-284. 

1954b. New names for two genera of Octocorallia. /. Wash. Acad. Sci. 44 (9) : 296. 

1956. Octocorallia. In, Treatise on Invertebrate Paleontology (Edited by R. C. Moore), 

Part F. Coelenterata, pp. 166-231. Geol. Soc. of America, New York. 

Brundin, J. A. F. 1896. Alcyonarien aus der Sammlung des zoologischen Museums in 

Uppsala. Bih. K. Svenska Akad. Vet.-Akad. Handl. 22 (4) : 1-22, pis. 1-2. 
Burchardt, E. 1902. Alcyonaceen von Thursday Island (Torres-Strasse) und von Amboina. 

II. Denkschr. med.-naturw. Ges. Jena, 8 : 653-682. 
Chalmers, D. 1929. The alcyonarian genus Siphonogorgia, with descriptions of new species. 

Proc. Roy. Phys. Soc. Edin. 21 : 159-169. (Also published in Thomson & Dean, 1931, 

pp. 149-152, 166-178.) 
Danielssen, D. C. 1887. Alcyonida. The Norwegian North- Atlantic Exped. 1876-1878, 

Zoology, Pp. viii -f- 169, 23 pis. Christiania. 
Deichmann, Elizabeth. 1936. The Alcyonaria of the western part of the Atlantic Ocean. 

Mem. Harv. Mus. Comp. Zool. 53 : 1-3 17, pis. 1-37. 
Gray, J. E. 1835. Characters of a new genus of corals (Nidalia). Proc. zool. Soc. Lond. 3 : 

59-60. 
1862. Description of two new genera of zoophytes (Solenocaulon and Bellonella) discovered 

on the north coast of Australia by Mr. Rayner. Ibid. 1862 : 34-37. 

1869. Notes on the fleshy alcyonoid corals (Alcyonium, Linn., or Zoophytaria carnosa). 

Ann. Mag. Nat. Hist. (4) 3 : 1 17-13 1. 

Harrison, R. M. 1909. On some new Alcyonaria from the Indian and Pacific Oceans, with a 

discussion of the genera Spongodes, Siphonogorgia, Chironephthya, and Solenocaulon. Trans. 

Linn. Soc. Lond. Ser. II, Zool. 11 (2) : 17-44, P^ s - 3~7- 
Hickson, Sydney J. 1903. The Alcyonaria of the Maldives. Part I. The genera Xenia, Telesto, 

Spongodes, . . . etc. In, Gardiner's The Fauna and Geography of the Maldive and Lacca- 

dive Archipelagoes, 2 (1) : 473-502, pis. 26-27. Cambridge. 

1930. On the classification of the Alcyonaria. Proc. zool. Soc. Lond. 1930 : 229-252. 

Kent, W. Saville. 1870. On two new genera of alcyonoid corals, taken in the recent expedition 

of the yacht Noma off the coast of Spain and Portugal. Quart. J. micr. Sci., n.s. 10 : 397- 

399, pi. 21. 
Klunzinger, C. B. 1877. Die Korallthiere des rothen Meeres. Erster Theil : Die Alcyonarien 

und Malacodermen. Pp. vii -f- 94, 7 pis. Berlin. 
Koch, G. von. 1891. Die Alcyonacea des Golfes von Neapel. Mitt. zool. Stat. Neapel, 9 (4) : 

652-676. 
Koch, W. 1886. Neue Anthozoen aus dem Golf von Guinea. Pp. (iv), 36, 5 pis. Marburg. 
Kolliker, Albert. 1875. Die pennatulide Umbellula und zwei neue Typen der Alcyonarien. 

Festschr. 25-jdhrigen Best. Phys.-Med. Ges. Wurzburg, 23 pp., 2 pis. 
Kukenthal, Walter. 1896. Alcyonaceen von Ternate. Nephthyidae Verrill und Siphono- 

gorgiidae Kolliker. Abh. Senckenb. naturf. Ges. 23 (1) : 79-144, pis. 5-8. 



120 A REVISION OF THE GENERA NIDALIA AND BELLONELLA 

Kukenthal, Walter. 1906a. Alcyonacea. Wiss.Ergebn. " Valdivia ", 13, i : i-iii, 12 pis. Jena. 

19066. Japanische Alcyonaceen. Abh. bayer. Akad. Wiss., Math.-Phys. Klasse, Suppl. 1 (1) 

1-86, 5 pis. Miinchen. 

1907. Versuch einer Revision der Alcyonarien. II. Die Familie der Nephthyiden. 3 Teil. 

Die Gattungen Eunephthya Verrill und Gersemia Marenzeller. Zool. Jb. (Syst.) 29 (5) : 
317-390. 

1910. Alcyonaria. 1 Teil. Fauna Siidwest-Aust. 3 (1) : 1-108, pis. 1-4. Jena. 

Macfadyen, L. M. I. 1936. Alcyonaria (Stolonifera, Alcyonacea, Telestacea and Gorgonacea). 

Sci. Rep. Gr. Barrier Reef Exped. 1928-29, 5 (2) : 19-71, pis. 1-5. 
Madsen, F. Jensenius. 1944. Octocorallia (Stolonifera-Telestacea-Xeniidea-Alcyonacea- 

Gorgonacea). Danish Ingolf -Exped. 5 (13) : 1-65, pi. 1. 
May, W. 1900. Beitrage zur Systematik und Chorologie der Alcyonaceen. Jena. Z. Naturw. 

33 for 1899 : 1-180, pis. 1-5. 
Molander, Arvid R. 1915. Northern and arctic invertebrates in the collection of the Swedish 

State Museum, VII. Alcyonacea. K. Svenska Vet.- Akad. Handl. 51 (n) : 1-94, pis. 1-3. 

1929. Die Octactiniarien. Further Zool. Res. Swed. Antarctic Exped. 1 901-1903, 2 (2) : 

i-iv, 1-86, pis. 1-5. 

Nutting, C. C. 191 2. Descriptions of the Alcyonaria collected by the U.S. Fisheries steamer 

" Albatross," mainly in Japanese waters, during 1906. Proc. U.S. Nat. Mus. 43 : 1-104, 

21 pis. 
Pfeffer, G. 1887. Ueber die Alcyonideen-Gattungen Nidalia, Gray, und Itephitrus, Koch. 

Verh. Ver. Hamburg, 6 : 101-104. (Not seen.) 
1889. Zur Fauna von Siid-Georgien. Jahrb. Hamburg. Wiss. Anstalt, Jahrg. 6 (2) : 49-55. 

(Not seen.) 
Putter, August. 1900. Alcyonaceen des Breslauer Museums. Zool. Jb. (Syst.), 13 (5) : 443- 

462, pis. 29-30. 
Simpson, J. J. 1905. Agaricoides, a new type of siphonogorgid alcyonarian. Zool. Anz. 29 : 

263-271. (Also published in Thomson & Henderson, 1906, pp. 15-19, pi. 10.) 
1907. On a new siphonogorgid genus Cactogorgia ; with descriptions of three new species. 

Trans. Roy. Soc. Edinb. 45 (3) : 829-836, 1 pi. (Also published in Thomson & Simpson, 1909, 

pp. 143-150, pl. 7)- 

1910. On a new species of Cactogorgia. Proc. Roy. Soc. Edinb. 30 (4) : 324-326, 1 pl. 

Stiasny, G. 1941. Alcyonaria und Gorgonaria aus dem Golf von Neapel. Pubbl. Staz. Zool. 

Napoli, 19 (1) : 1-47. 
Strand, E. 1928. Miscellanea nomenclatorica zoologica et palaeontologica. Arch. Naturg. 

92 (A8) : 31-36. 
Studer, Th. 1878. Uebersicht der Anthozoa Alcyonaria, wahrend der Reise S. M. S. Gazelle 

um die Erde gesammelt wurden. Mber. preuss. Akad. Wiss. 1878 : 632-688, pis. 1-5. 
1890. Note preliminaire sur les Alcyonaires provennat des campagnes du yacht l'Hirondelle 

(1886-1887-1888). Mem. Soc. zool. Fr. 4 (2) : 86-95. 
1901. Alcyonaires provennant des campagnes de l'Hirondelle (1886-1888). Result. Camp. 

Sci. Monaco, Fasc. 20 : 1-64, 11 pis. Monaco. 
Thomson, J. Arthur. 1927. Alcyonaires provenant des campagnes scientifiques du Prince 

Albert i er de Monaco. Ibid., Fasc. 73 : 1-77, 6 pis. Monaco. 
Thomson, J. Arthur & Dean, Laura M. I. 1931. The Alcyonacea of the Siboga Expedition, 

with an addendum to the Gorgonacea. Siboga-Exped. 13d : 1-227, pl s - 1-28. Leiden. 
Thomson, J. Arthur & Henderson, W. D. 1905. Report on the Alcyonaria collected by 

Professor Herdman at Ceylon in 1902. Ceylon Pearl Oyster Fisheries-1905- Suppl. Reports, 

No. 20 : 269-328, pis. 1-6. 
1906. An account of the Alcyonarians collected by the Royal Indian Marine Survey ship 

" Investigator " in the Indian Ocean. I. The Alcyonarians of the deep sea. Pp. xvi + 132, 

10 pis. Calcutta. 
Thomson, J. Arthur & Mackinnon, Doris L. 1910. Alcyonarians collected on the Percy 

Sladen Trust Expedition by Mr. J. Stanley Gardiner, M.A., F.R.S. Part 2. The Stolonifera, 

Alcyonacea, Pseudaxonia and Stelechotokea. Trans. Linn. Soc. Lond. Zool. ser. 2, 13 (8) : 

168-21 1, pis. 6-14. 



A REVISION OF THE GENERA NIDALIA AND BELLONELLA 121 

Thomson, J. Arthur & Rennet, Nita I. 1927. Report on Japanese pennatulids. /. Fac. Sci. 

Imp. Univ. Tokyo, Sect. IV, Zool. 1 (2) : 1 15-143, pis. 7-9. 
Thomson, J. Arthur & Simpson, James J. 1909. An account of the Alcyonarians collected by the 

Royal Indian Marine Survey ship " Investigator " in the Indian Ocean. II. The Alcyonarians 

of the littoral area. Pp. xii -{- 319, 9 pis. Calcutta. 
Thomson, J. Stuart. 1910. The Alcyonaria of the Cape of Good Hope and Natal. Alcyonaria. 

Trans. Roy. Soc. Edinb. 47 (3) : 549-589, pis. 1-4. 

192 1. South African Alcyonacea. Trans, roy. Soc. S. Afr. 9 (2) : 149-175, pis. 5-6. 

Tixier-Durivault, A. 1954. Les octocoralliaires d'Afrique du Sud (I. Alcyonacea). Bull. 

Mus. Hist, nat., Paris, (2) 26 : 124-129, 261-268, 385-390. 
Utinomi, Huzio. 1954. Some Alcyoniid octocorals from Kii coast, middle Japan. Publ. seto 

mar. biol. Lab. 4 (1) : 43-55, pi. 1. 
1957. The alcyonarian genus Bellonella from Japan, with descriptions of two new species. 

Ibid. 6 (2) : 147-168, pis. 9-10. 
Wright, E. P. & Studer, Th. 1889. Alcyonaria. Rep. sci. Res. " Challenger " Exped. 31, 

pt. 64. Pp. lxvii -f- 314, 49 pis. London. 
Yamada, Mayumi. 1950. Descriptions of two Alcyonium from Northern Japan. Annot. zool. 

Japon. 23 (3) : 114-116. 




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EAR PLUG LAMINATIONS IN 

RELATION TO THE AGE 

COMPOSITION OF A POPULATION 

OF FIN WHALES 

(BALAENOPTERA PHYSALUS) 



P. E. PURVES 

AND 

M. D. MOUNTFORD 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 6 

LONDON: 1959 



EAR PLUG LAMINATIONS IN RELATION TO 
THE AGE COMPOSITION OF A POPULATION 

OF FIN WHALES 
(BALAENOPTERA PHY S ALUS) 



BY 

P. E. PURVES 

British Museum (Natural History) 
AND 

M. D. MOUNTFORD 

The Nature Conservancy 



Pp. 123-161, Plates 5-6 ; 10 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 6 

LONDON: 1959 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5, No. 6 of the Zoological series. 




Trustee of the British Museum. 1959 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued January, 1959 Price One Pound 



EAR PLUG LAMINATIONS IN RELATION TO 
THE AGE COMPOSITION OF A POPULATION 

OF FIN WHALES 
(BALAENOPTERA PHYSALUS) 

By P. E. PURVES and M. D. MOUNTFORD 

SYNOPSIS 

i . The method of grinding the ear plugs of Fin Whales and of counting their laminae is briefly 
described. 

2. The growth of the plug is compared with that of the skull width and body length. 

3. A provisional rate of formation of the laminae is assessed and correlated with previously 
established data about the growth and age of Fin Whales. 

4. From the age frequency distribution based on the ear plug lamination analyses the 
apparent natural mortality rate for the " Sanctuary " population is demonstrated. 

In the course of a detailed description of the ear plug of the Mysticeti (Purves, 1955) 
it was suggested that there might be a correlation between the laminar structure 
of the core of the plug and the age of the animal from which it was taken. Further 
support was given to this hypothesis by the examination of 18 Fin Whales at 
Steinshamn, Norway, during the summer of 1955, when the age as estimated from 
the ear plugs was compared with that assessed from the ovaries and baleen plates, 
Laws & Purves (1956), Ruud (1945). 

The results of this enquiry were sufficiently encouraging to warrant further investi- 
gation. Since the National Institute of Oceanography is concerned with age deter- 
minations, especially in studies of the life cycle and populations of whales, Dr. 
Mackintosh and Dr. Laws had made arrangements, after the publication of the 
paper by Purves (1955) for the collection of a large number of ear plugs from whales 
taken by factory ships in the Antarctic season of 1955-56. The plugs were obtained 
from some 454 whales, together with relevant data on the whales from which they 
were taken. Before examining the material themselves however, they were good 
enough to make the collection available to us so that the relation between the lami- 
nations of the plug and the age of the whales could be further examined. This is 
the subject of the greater part of the present paper. It was a matter of much 
interest however, to examine the age composition of the population from which the 
plugs were obtained and by agreement with the National Institute of Oceanography 
the paper has been extended beyond its initial scope to include this aspect. 

zool. 5, 6. 6 



126 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

MATERIAL AND METHOD 

The ear plugs had been wrapped singly or in pairs and preserved in 5% formalde- 
hyde on board the factory ships and were consequently hard and in good condition 
for examination in the laboratory. It is unfortunate that a number of plugs had 
become detached from their wrappings during transit as a result of which it has 
been impossible to determine the length and sex of the animals involved, but even 
these specimens have been useful in plotting the age frequency distribution. 

Because of the asymmetry and extreme attenuation of the distal end of the core 
it was thought that bisection of the plug would result in loss or damage to the earliest 
formed part of the structure, so each specimen was carefully group down to the 
central plane by hand. The grinding was carried out by rubbing the plug with a 
rotary movement against waterproof abrasive cloth which had been cemented with 
rubber solution to a sheet of plate glass. Coarse and fine grades of abrasive were 
used and during the whole operation a stream of water was directed over the surface 
of the plate. It is appreciated that for routine examination of the plugs a mechanical 
grinding plate or stone would be preferable, but since the time expended on each 
plug using the more primitive method amounted to no more than three minutes, 
the above described apparatus was considered adequate for the present sample. 

Plate 5 shows a series of ear plugs from female Fin Whales, the specimens being 
chosen at random from groups of plugs which differed consecutively in lamination 
number by four laminations. The series shows a progressive lengthening of the core 
of the plug and a gradual darkening of its matrix from the external shell towards 
the central axis. From the plate it appears that the overall length of the core of 
the plug is no guide to the number of laminations which it contains, but it will be 
shown later that in spite of the great variability in the lengths of the cores of equal 
laminar number, the average length of the core per lamination number is correlated 
with the number of laminations. It may be stated that on the whole, the diminution 
in the thickness of the laminations from the distal to the proximal end of the core 
is more regular in plugs from males than in those from females. This characteristic 
is not always apparent in a small sample, and cannot be used as a guide to the sex 
of a whale from which any one plug originated but may be of use statistically in 
connection with a large sample of specimens. The unwrapped specimens referred 
to on page 161 were divided on the basis of this feature but the information so obtained 
has been of limited use and is referred to with reservation in the present paper. 

In the plugs from immature and very young animals the primary laminations may 
be subdivided into a number of ill-defined, subsidiary layers but the latter become 
obliterated as more primary laminae are formed and it has been the practice through- 
out this investigation to treat every lamination in the older specimens as a single 
unit, however narrow and apparently subsidiary In the very old specimens the 
proximal end of the core may appear to the naked eye or with dissecting binoculars 
to be quite undifferentiated, but when examined microscopically, these undifferentia- 
ted areas are seen to be made up of a series of regularly-spaced refractive layers. 
When these layers are counted towards the distal end of the core they are observed 
to increase gradually in thickness and in the latter respect to be in geometrically 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 127 

progressive sequence with the coarser layers which are visible to the naked eye. 
The microscopic section shown in Plate 6 was taken from the proximal end of the 
only plug in which for some unknown reason there was a sharp transition from very 
coarse to very fine laminae, but it is useful in that a comparison can be made between 
the two types of laminae within the compass of a very small section. The section 
shows the appearance of the refractive layers referred to above after removal of 
the ceruminous component. The area bottom left, which shows the lateral extremity 
of the short axis of the base of the core originally contained very little cerumen 
and consists of a mass of undifferentiated squamae of keratin. On the right, the 
squamae are broken into a series of laminae of approximately equal thickness, 
each of which contains a number of flattened nuclei. 

The upper part of the section shows two of the very coarse laminae. When 
stained with haemalum and Mallory's triple stain the greater part of the keratinized 
mass appeared bright blue, but in each lamina there was a conspicuous band of 
orange which occupied a position immediately distal to the layer of flattened nuclei. 
These bands of orange, which are deemed to mark areas of imperfectly keratinized 
cellular matter accompanying degenerate nuclei, seem to be the main distinguishing 
feature of the laminations and each can be used as a criterion of what is a single 
lamination. If it were possible to cut and stain the whole core in this manner 
the lamination number could be estimated with great accuracy. In the more simple 
method of counting the laminations on the unstained cut surface of the core it 
would be improbable to make an error of more than plus or minus 4 laminae in 
each plug and with a large sample, such as the one under consideration, errors of 
this magnitude would probably cancel each other out. 

All the specimens referred to in this paper were collected aboard the factory 
ships Baleana, Southern Harvester and Southern Venturer. The Southern Harvester 
collection was obtained from that sector of the Antarctic known as the " Sanctuary " 
(Lat. 6o° W.-120 W.) and is referred to as the Area I sample. The Balaena 
collection was obtained from Area II whilst the Southern Venturer collection was 
obtained from both sectors. The latter collection has been broken up and amal- 
gamated with the Balaena and Southern Harvester specimens so that only the two 
populations, the Area I and Area II samples are described. 

The data referred to in the text are given in Tables A to D of the Appendix. 

From the results of the examination of the Steinsham material (loc. cit.) it has 
been established that there is a direct correlation between the number of laminations 
in the core of the plug and the age of the animal as assessed from the ridges on the 
baleen plates. This correlation can however be shown only in respect of animals 
under 6 years of age, since the analysis of the baleen plate data becomes difficult 
after this age. 

Before attempting to obtain the age frequency distribution it is necessary to 
establish whether or not this age lamination correlation continues throughout life. 
It may reasonably be assumed that the growth of cetaceans follows a pattern similar 
to that which is found in most other mammals and that there is an age shortly 
after sexual maturity beyond which further increases in bodily proportions are small 
relative to the immature growth increments. This being so, one might expect a 

zool. 5, 6. 6§ 



128 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

certain variability in the adult dimensions due to the differential immature growth 
rates. If a simple relationship can be shown between the skull width and the lami- 
nation number as suggested by Purves (1955) then the lamination number would 
be valueless as a means of estimating the individual ages of a variable adult population 
the individual growth rates of which are unknown. 

Unfortunately no data relevant to skull width are available for the present sample 
so that the skull-width-lamination number relationship cannot be found directly. 
Mackintosh & Wheeler (1929) have published a very comprehensive list of skull 
widths and body lengths of Fin Whales so we may use their figures relevant to the 
range of lengths available in the present sample to find the body-length skull-width 
relationship. 

The Body Length, Skull-width Relationship 

In the Tables of the External Characters of Fin Whales, Mackintosh & Wheeler 
(1929) quote the skull widths and total lengths of 162 female and 206 male Fin 
Whales taken at South Georgia during the years 1926 and 1927. The hypothesis 



33 

32 




















FEMALES 






























3/ 


























30 


























29 


























26 
























• • 


27 


























26 






















• • • . . -* 

•• 

: • % 


^^» ' 


2-5 






















. .. .Xj^ 


• . 


24 
23 




















• 


•••f&* » • • • 


• 


11 
















• 


• 




• • 

• ♦ • • • 

• 

•• 




11 








• 




• 






» 


• 






20 










• 




• 


* 


• 


• 


• 




1-9 




• 


• 




• 


• 


• 


• 










18 


• 




• 


<f 


* 


•• 




• 


• 









10 



zz 



23 



IS 



Length of body (in metres) 

Fig. 1 . The skull width-body length relationship of a population of female Fin Whales 
Balaenoptera physalus measured at South Georgia, Mackintosh & Wheeler (1926-27). 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 129 

of a linear relationship between the skull-width and total length provides an excellent 
fit to the data. The linear regression of skull width on total length (Text-figs. 
1 and 2) is found to be : 

y = 0-14029 -f 0-10209* for females (Text-fig. 1) and y = 0-02946 + 0-10813% 
for males (Text-fig. 2) where y and x are the skull width and total length in metres 









MALES 




*•? 














26 








• • 






ZS 








• • • 
• # • 






Z-4 
23 

22 
Zl 
20 
19 






• 
1/ 


» • • %^^^ 

* • ^>a • • 
• ••«•*•• **"&** •* * 

• • 1 *• ^z •• •••• • # * •• 

^^ • • • 






18 


• 


• ^* 


• 


• 






'7 


v* 


<£• 


• 








IS 




• 










14 














IS 






























fS 14 IS 


lb 


a 


/tf /$ 20 2/ 22 


2 


J 



Length of body (in metres) 

Fig. 2. The skull width-body length relationship of a population of male Fin Whales 
Balaenoptera physalus measured at South Georgia, Mackintosh & Wheeler (1926-27). 



respectively. It must be emphasized that the linear relationship is only valid in 
the above range. It is of incidental interest to note that the rates of increase of 
skull width on total length for female and male whales are not significantly different. 
A measure of the high degree to which the association between skull width and 
total length approaches a linear relationship is given by the correlation coefficient 
which is as great as 0-87 for the females and o-8o for the males. There is thus no 
justification for rejecting the hypothesis of an isometric relationship between the 
skull width and total length of the Mackintosh & Wheeler population. It is a 
justifiable inference to conclude that this isometric relationship holds for all popu- 
lations of this species in which the total lengths lie in the range 14 metres to 24-3 



130 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

metres. In particular it holds for populations in Areas I and II. It is therefore 
possible to deduce the lamination-number skull-width relationship from that holding 
between the lamination number and total length. 

It is of some bearing on the above topic to compare the Mackintosh & Wheeler 
sample with the Area I sample for their common measurement of total length. 
The former population is truncated of several extreme small values so that the bottom 
limit for each sex in the two samples is the same. It has been necessary for ease of 
comparison to convert Mackintosh & Wheeler's small i-metre length ranges to the 
mean value in feet. The figures are given below : 



Male 


Fin Whale 


Female Fin Whales 


S. 


Georgia 


S. 


Georgia 




jl 




JL 


t 


■\ 


r 


1 




Number of 




Number of 


Length 


measurements 


Length 


measurements 


55 


6 


— 


— 


58 


7 


58 


6 


61 


13 


61 


11 


64 


45 


64 


9 


67 


78 


67 


30 


70 


45 


70 


36 


73 


3 


73 


38 


77 


1 


77 


8 


80 


— 


80 


1 


ncy distribution of th 


Le specimens 


from Area I 


Male Fin Whales 


Female Fin Whales 


Area I 


Area I 




a 




a 


r 


^ 


r 


^ 




Number of 




Number of 


Length 


specimens 


Length 


specimens 


55 


3 


— 


— 


58 


3 


58 


5 


61 


12 


61 


12 


64 


49 


64 


17 


67 


76 


67 


30 


70 


26 


70 


60 


73 


2 


73 


30 


77 





77 


7 


80 


— 


80 


1 



given below 



Inspection of the above lists of measurements shows that the length frequencies 
of both samples are arranged more or less symmetrically about the mean lengths. 
The mean lengths and standard deviation for the two samples were found to be : 







Mean length 


Standard deviation 


S. Georgia 


Males 


66-o 


3-6 


Area I . 


,, 


66-o 


3*1 


S. Georgia 


Females 


68-7 


4'6 


Area I . 


,, 


68-6 


3'3 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 131 

It is obvious that the two samples are remarkably alike in their distribution of the 
total lengths. On the other hand the 38 female and 23 male specimens caught in 
Area II give the following value : 

Mean length Standard deviation 

Area II . . Females . 70*1 . 4-5 

Area II . . Males . 67-0 . 3-7 

Although as shown by a " t test " the differences in mean total length between the 
Area II and the Area I whales are not significant at the 95% level there is some slight 
evidence for suspecting that the mean total lengths of both sexes in Area II are 
approximately 1 ft. greater than those in the Area I sample. The values of mean 
total length given above are only valid over the length range 55 ft. to 80 ft. and would 
be rather lower over the whole length range of the population. 

The Lamination Number-Body-length Relationship 
In Text-figs. 3 and 4 the lengths of males and females from Areas I and II respec- 
tively have been plotted against the lamination number. Certain facts which will 
be examined in detail are given below : 

(a) There is considerable variation in length at every lamination number. 

(b) At every lamination number the average length of the females is greater 
than that of the males. 

(c) The average length per lamination number of the specimens from Area II 
is greater than that of the specimens from Area I. 

(d) After the formation of the eighth lamination the number of individuals 
in each laminar group is inversely related to the number of laminations. 

The mean body length for each lamination group is given in the following tables : 





Sample Area 1 








Female Fin Whales 


Male Fin Whales 




,A 




^ 






r 


^ 


r 


A 




Number 


Mean 


Number 


Mean 


amination 


of 


length 


of 


length 


number 


specimens 


in ft. 


specimens 


in ft. 


3* 


— 


— 


1 


55 


4 


— 


— 


2 


69 


5 


1 


58 


— 


— 


6 


4 


65-75 


1 


64 


7 


8 


64 


2 


58 


8 


6 


61 


2 


64 


9 


11 


64 


8 


63-4 


10 


9 


65 


5 


63-4 


11 


6 


65 


3 


63 


12 


8 


67-7 


5 


64 


13 


3 


67 


8 


66 


14 


7 


69 


9 


65-3 


*5 


3 


70 


5 


66 


16 


8 


72 


9 


66 


17 


3 


7i 


7 


67 


18 


4 


71 


1 


69 



132 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 





Sample 


Area I — c 


ont. 






Female Fin Whales 


Male Fin Whales 




a. 




a 






r 


■\ 


r 


-\ 




Number 


Mean 


Number 


Mean 


Lamination 


of 


length 


of 


length 


number 


specimens 


in ft. 


specimens 


in ft. 


19 


1 


74 


2 


66 


20 


5 


7i 


5 


66 


21 


1 


74 


2 


70 


22 


4 


7i 


5 


68 


23 


1 


73 


7 


67 


24 


8 


72 


5 


67 


25 


2 


7i 


3 


66-6 


26 


6 


72 


3 


67 


27 


3 


72 


2 


65 


28 


2 


7i 


3 


66-3 


29 


2 


68-5 


2 


69 


30 


1 


73 


4 


66-5 


3i 


1 


70 


2 


66-5 


32 


5 


74 


1 


65 


33 


1 


75 


1 


70 


34 


2 


— 


1 


7i 


35 


1 


75 


4 


66 


36 


6 


72 


1 


68.5 


37 


1 


76 


1 


65 


38 


— 


— 


1 


67 


39 


1 


73 


1 


69 


40 

4i 
42 


5 


7i 


2 


68-5 


2 


73 


1 


66 


43 


1 


69 


3 


66-5 


44 


— 


— 


2 


66 


45 


1 


72 


1 


68 


46 


1 


75 


2 


67 


49 


— 


— 


1 


67 


50 


2 


75 


3 


67 


52 


— 


— 


2 


66 


53 


— 


— 


1 


65 


54 


2 


76 


3 


66 


55 


1 


69 


1 


66 


56 


3 


7i 


2 


64 


58 


— 


— 


2 


66-5 


59 


— 


— 


1 


67 


60 


— 


— 


2 


69 


61 


— 


— 


1 


68 


62 


— 


— 


1 


73 


63 


— 


— 


1 


70 


66 


— 


— 


1 


65 


67 


— 


— 


1 


70 


70 


1 


75 


1 


7i 


7i 


— 


— 


1 


65 


75 


— 


— 


1 


68 


76 


— 


— 


2 


66-5 


80 


1 


73 


— 


— 


83 


— 


— 


1 


64 


85 


— 


— 


I 


64 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 133 



Sample Area II 





Female Fin Whales 


Male Fin Whales 




■*- 




^ 






r 


"\ 


r 


% 




Number 


Mean 


Number 


Mean 


Lamination 


of 


length 


of 


length 


number 


specimens 


in ft. 


specimens 


in ft. 


3 


1 


60 


— 


— 


4 
5 


2 


64*5 


1 


55 


6 


1 


63 


— 


— 


7 


3 


61 


— 


— 


8 


1 


74 


2 


63 


9 


3 


65-5 


4 


64 


10 


3 


67 


1 


63 


11 


3 


72 


1 


66 


12 


5 


70-6 


— 


— 


13 




72-0 


— 


— 


14 




72 


1 


65 


15 




72 


— 


— 


16 




68 


2 


69*5 


17 


— 


— 


1 


68 


18 




76 


— 


— 


19 




76 


1 


71 


20 




72 


2 


71 


21 




73'6 


— 


— 


22 




72-5 


1 


70 


23 


— 


— 


2 


69 


24 




75 


— 


— 


28 




73 


— 


— 


29 




77 


— 


— 


30 




75 


1 


69 


31 




73 


1 


70 


32 




75 


— 


72 


33 




75 


— 


— 


34 




75 


2 


68 


26 


— 


— 


1 


70 


38 




68 


— 




40 


— 


— 


2 


69 


42 




72 


— 


— 


43 




75 


1 


70 


44 




78 


1 


7i 


45 


— 


— 


1 


67 


46 




77 


— 


— 


47 




74 


— 


— 


48 




76 


— 


— 


50 




74 


1 


67 


52 




75 


— 


— 


53 




79 


— 


— 


59 


— 


— 


1 


65 


62 




75 


— 


— 


63 


— 


— 


1 


76 


66 




78 


— 


— 


7 6 


— 




— 


63 



134 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 





















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EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 135 

Text-figs. 3 and 4 suggest that for each sex, whales with more than 13 laminations 
are such that their total lengths are almost independent of their lamination numbers, 
i.e. the correlation coefficient between lamination number and length is not greatly 
different from zero for whales with more than 13 laminations in the ear plug. 
This correlation coefficient was evaluated as 0-169 for females and 0-27 for males 

in Area I. Using t = r / ^ n ~ ' as a if-variate where " r " and " n" are the corre- 
8 V (1-r 2 ) 

lation coefficient and the number of individuals in the sample respectively, both of 
the above coefficients were found to be not significantly different from zero. This 
same analysis applied to the Area II whales revealed that as in Area I the lamination 
number was nearly independent of total length for whales with more than 13 lami- 
nations, i.e. for any Fin Whale with a lamination number greater than 13 the specific 
value of its number can be considered to have little bearing on its total length and 
skull width. 

If any one plug is measured from its base to each successive lamination the increase 
in core length is found to take the form of an exponential curve like that which was 
described by Purves (1955), but the total length of the core for any one lamination 
number varies very greatly from one specimen to another, and cannot be used even 
as a rough guide to the lamination number. An average growth curve has been 
obtained (Text-fig. 5) by plotting the mean length of the core per lamination number 
at every fourth lamination against the lamination number. These mean core 
lengths are remarkably similar in the two sexes notwithstanding the difference of over 
4 ft. in the adult body lengths between males and females. The curve appears to 
show that plugs of high lamination number are disproportionately long, but this 
effect may be due to the small number of plugs of high lamination number in the 
sample. If high lamination number is an indication of old age the presence of these 
disproportionately long plugs may be due to the mixing of populations owing to 
lateral migration Brown (1954). It will be seen at once that this growth curve is in 
no way comparable with that of skull width against body length so that neither 
the lamination number nor the plug length have any close relationship to the body 
length and skull width. It must, therefore, be concluded that the growth of the 
plug is independent of that of the body as a whole, and that its laminated core forms 
part of a rhythmic growth system which was initially established in response to 
one or more of a number of factors involving the passage of time, such as the breeding 
cycle, migratory movements, nutritional changes, temperature variation or inherent 
mammalian moulting cycles. The periodicity of the rhythm may vary from one 
individual to another but in a population with a very regular cycle of behaviour the 
variation is likely to be very small. Since the growth of the plug is not conditioned 
by the skull dimensions, it is possible that the shape of the bony meatus is continually 
adjusted to the growth increments, that there is no resorption of any part of the 
plug, and that the laminations constitute a complete record of the periodic des- 
quamations from birth to death. In this connection it is of interest to draw attention 
to the difference between the growth of the ear plug in whales and that of the cycloid 
and ctenoid scales of fishes. Van Oosten (1955) states that in the fishes cessation 
of body growth ultimately ends in suspensions of scale formation. He continues 
zool. v, 6. 6SS 



136 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

" In maintaining coverage of the body the scales have been found to grow at an 
approximately fixed ratio with the fish. The growth of the scale, is therefore, more 
or less a replica of that of the body. This fact has made possible the employment of 
scales in estimating the past growth of an individual. Multiplying the ratio of the 
length of that part of the scale that was completed at the end of a certain year of 
life to the final length of the scale by the length of the fish gives the estimated length 




Fig. 5. Lamination number-core length relationship of the ear-plugs of 
a sample of Fin whales Balaenoptera physalus. 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 137 

that the individual attained at that particular age. Yearly increments of growth 
can be obtained by subtracting the different computed lengths ". A similar system 
of length estimation based on the thickness of ear plug laminae could be arrived at 
with some difficulty in respect of whales under 6 years of age, but for animals above 
this age no correlation is possible. Van Oosten also states that age determination 
in fishes permits studies on variation in growth rate with species, latitudes and differ- 
ent bodies of water. " By observing the time of formation of laminae the length of 
growing seasons may be determined." It is very doubtful whether environmental 
conditions have any direct influence on the time of formation of the laminae of the 
ear plug in whales, although nutritional conditions may to some extent determine 
their thickness. Since there is strong evidence that lamina formation is an inherent 
moulting cycle it is very probable that it takes place whether the whale migrates or 
not. If the rate of moulting can be established the ear plug would probably be a 
more accurate age indicator than the fish scale. 

Since it was shown by Laws & Purves (1956) that up to the formation of the 12th 
lamination a fairly close correlation exists between body length and lamination 
number and in the present paper that after the 13th lamination the correlation 
coefficient approaches, but is not precisely, zero, it might be expected that an expo- 
nential growth function could be demonstrated over the whole range of laminations. 

In Text-fig. 3 curves of mean total length against lamination number have been 
fitted by the method of least squares to the samples of males and females from 
Area I. Dealing first with Area I females it was found that if " y " (in feet) is the 
length and " x" is the lamination number then a good fit to the data is given by 

y = 72-91 — 24-24 exp (— 0-1253;) 

where " exp " is the exponential function. This expression is a description of the 
data only over the lamination range 6-80 and gives a growth curve from 61-5 ft. 
at 6 laminations to a maximum length of 72-91 ft. ; extrapolation beyond these limits 
is not justified but Mackintosh & Wheelers' (1929) curve of immature body lengths 
can be used to complete the curve. 1 It may be noted that a length of 71-9 ft. is 
attained at the formation of the 26th lamination. The line in the graph (Text-fig. 3) 
represents the growth function, which is such that at any given lamination number 
the growth rate is proportional to the " remaining size " i.e., the maximum size 
minus the size reached. A similar growth curve was fitted to the males Area I 
(Text-fig. 3) and a good fit was obtained by the curve 

y = 67-31 — 19-55 exp (— O'iBx). 

Here again the curve only describes the growth in the range 6-80 laminations. 
The estimated length of 60-7 ft. at 6 laminations increases to an average maximum 
of 67-31 ft. The length of 66-3 ft., one foot less than the maximum is achieved at 
about the 16th lamination. The points marked on the 2 diagrams are the mean 
values of the lengths for each lamination number. This result is in accordance with 
previously established information regarding the difference in the growth rates 

1 Because of the under-representation in the sample of the 6th lamination group the first part of the 
growth curve may give an over-estimation of the average body length in this group, see page 149. 



138 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

between male and female Fin Whales and both curves approximate to the average 
mammalian growth curve. It will, therefore, be assumed that the laminations are 
laid down at fairly regular intervals throughout the life of the animal. 



The Rate of Formation of Laminae 

In order to establish the rate of formation of the laminae, use can be made of the 
following information : 

(a) The lowest number of laminae formed when the majority of the females 
are either lactating or pregnant. 

(b) The lowest number of laminae formed when the vertebral epiphyses are 
ankylosed to the centra in the majority of adults. 

The difference between these two figures will represent the approximate number of 
laminae formed between sexual and physical maturity which by deduction from 
Wheeler's (1929) figure of 15 corpora lutea accumulated in the ovary and Laws' 
(1955) rate of 1-4 per annum for their accumulation should be approximately 10 years. 

In Text-figs. 6 and 7 each horizontal line represents the vertebral column and can 
be considered to be divided into four sections containing the anterior thoracic, 
posterior thoracic, lumbar and caudal vertebrae respectively. Each vertebral 
column or portion of a vertebral column has been placed in one or more of the vertical 
divisions according to the state of fusion of the vertebrae. A category described as 
" Fused Joint Visible " was given in the original data but for the purpose of the 
charts it was assumed that if the joint was visible it must also contain a thin layer 
of cartilage, so vertebrae in this category were placed in the division labelled " Unfused 
Fine Cartilage ". The horizontal lines in Text-fig. 6 have been thickened to denote 
whales which were known from the data to be pregnant. 

The horizontal lines which completely traverse the charts are drawn through 
the lamination numbers above which the vertebral epiphyses are fused and invisible 
in the majority of individuals, and through the number in the chart of the female 
whales above which the majority are pregnant. It will be seen from Text-fig. 6 
that the number of laminations shown between these two lines is 20 so that if the 
estimated period between sexual and physical maturity is 10 years the rate of lami- 
nation formation is approximately 2 per year. 

As a check on the above result it may be noted that Mackintosh & Wheeler's 
figure for the length of female Fin Whales at sexual maturity is 66 ft. This figure 
has been confirmed by Peters (1939), Mackintosh (1942), Brinkmann (1948), Nishiwaki 
& Hayashy (1950), the length at physical maturity being in the region of 72-4 ft. 
The regression line of growth (Text-fig. 3) passes through these two lengths at the 
10th and 30th lamination respectively. If the period between sexual and physical 
maturity is 10 years the rate of accumulation of laminae is 2 per year. When 
the figures of Nishiwaki & Oye (195 1) and Jonsgard (1952) of 67 ft. at sexual maturity 
and 75 ft. at physical maturity are applied to the growth curve of the Area II females 
(Text-fig. 4) they are seen to cut the curve at the 8th and 28th lamination respectively. 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 139 

Here again if the period between sexual maturity and physical maturity is 10 years 
then the rate of accumulation of laminae is 2 per year. 

Up to date no collection of Blue Whale ear plugs large enough for statistical 
analysis has been received but from inspection of the few specimens which were 
collected by Simons (1956) there is no reason to conclude that the rate of accumulation 
of laminae differs greatly from that estimated in the Fin Whale ear plugs. In 
this connection a letter received from A. H. Laurie is quoted below : 

' Your query as to the interpretation of the figures given in my paper (Laurie, 
1937) has resulted in the unearthing of an error which appears to have lain 
unnoticed for twenty years. On page 250 I have shown that the annual incre- 
ment of corpora lutea, now called corpora albicantia, appears to be slightly in 
excess of one per annum. The average figure given was 1-13. 

" As is seen in the tables, the above conclusion is based on samples containing 
three categories of whale, namely pregnant, resting (i.e. neither pregnant nor 
having recently ovulated), and non-pregnant but having recently ovulated. 

For convenience I reproduce here the totals in this argument : 







Percentage 


Percentage 


Percentage 


recently 


pregnant 


resting 


ovulated 


64 


24 


12 



" Where the mistake arises, and I have repeated it categorically in section 
6 of the Summary (p. 268) is that I have taken the annual increment to apply 
to the whole adult stock of female Blue Whales. At the same time no account 
has been taken of that fraction of the stock which was lactating, and thereby 
absent from the sample. To clarify this statement let us add a hypothetical 
but plausible number to the above percentages, to include the absent lactating 
whales and let us assume that the number of lactating whales corresponds to 
a similar percentage of pregnant whales in the previous year. The total stock 
is then represented by 100 (as above) plus an additional 64. 

" The total is thus 164, of which the additional 64 just added, being in lactation 
can be presumed not to have ovulated during the year under review. It 
follows therefore that the figure given for annual increment of corpora albicantia 
must be corrected thus : 

100 r 

1-13 x 164- = °' 69 - 

" Another way of stating the amendment is to say what I should have said 
in the first place, namely that the average increment in corpora lutea per 
breeding cycle is 1-13. 

" If we now employ the revised figure of 0-69 c.l. per annum for Blue females 
as a whole, the putative time scale can be revised as follows. 

" Physical maturity was shown to coincide with the accumulation of n 
corpora. After allowing 1-91 corpora for the first breeding season, i.e. to include 



140 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 



WHALE LAM. 
NO. NO. 


UNFUSED THICK 
CARTILAGE 


UNFUSED FINE 
CARTILAGE 


FUSED JOINTS 
INVISIBLE 


358 80 

700 70 

2009 66 

1352 56 

1703 55 
1250 54 
1435 50 

706 46 
438 45 
2130 43 
1042 42 
2085 42 
488 40 

1704 40 
937 40 

2015 40 

1044 40 

6670 38 

1931 37 

1669 36 

593 36 

2007 36 

1933 36 

741 36 

780 35 

482 34 

1633 36 

1234 32 

486 32 

1710 32 












































































672 32 

1280 30 

1968 30 

747 29 

1169 29 

743 28 

1704 27 

1625 27 

478 26 

1358 26 

867 26 

2099 26 

745 26 

749 25 

398 24 

434 24 

1366 24 

603 24 

700 24 

1892 24 

1788 24 

894 24 

1793 23 

714 22 

2122 22 

1779 22 

1083 22 

646 21 

597 20 

898 20 

1964 19 

1085 18 

2136 18 

869 18 

784 18 

1854 17 

737 17 

513 17 

1823 16 

703 16 

855 15 

1811 15 

1079 14 

906 14 

1784 14 

1968 14 

2128 14 

656 14 

670 13 

2094 12 

786 12 

1034 12 

1320 12 

448 12 

1208 12 

















^— 






































































"™~~~" 






— 
















































1167 12 








1 





EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 141 



1165 

935 

1817 

1821 

1890 

1937 

1196 

1279 

859 

1247 

1898 

1749 

519 

1813 

1036 

1248 

118 

933 

1108 

1748 

2126 

1930 

1864 

2059 

1454 

871 

1856 

776 

652 

707 

601 

648 

1819 

1785 

739 

1200 

2013 

1820 

352 

1714 

863 

1284 



Fig. 6. State of fusion of the vertebral epiphyses of a sample of female antarctic Fin whales 
Balaenoptera phy stilus. Each horizontal line represents a vertebral column which can be 



AT PT 



considered to be divided thns | • \ 
PT = posterior thoracic, L = lumbar, C 
which were known to be pregnant. 



I *» ! £ i where AT = anterior thoracic 
caudal. The thick lines represent individuals 



a period of 2 J years from birth, the remaining 9-1 corpora could represent 
13-2 years so that the age at physical maturity now becomes 2-5 + 13-2 = 15-7 
years, instead of 10 to n years. The oldest whale in my collection was approxi- 
mately 45 instead of 30. 

" I must emphasise that the above figures relate to the ' ovary clock ' of more 
than twenty years ago. In view of the much higher percentage of pregnancy 
now observed in whale samples, the regulator of the clock has apparently been 
altered, presumably by external influences." 

It will be noted that the age at physical maturity for Fin Whales was assessed at 
15 years and that the oldest specimen recorded in the present sample was 42 \ years 
old. From the ear plug and ovary data therefore, both Fin and Blue Whales appear 
to become physically mature at the same age and it may be presumed that they have 
much the same maximum life span. 

Regarding the male Fin Whales the evidence is rather less conclusive. According to 
Jonsgard (1952) " the various investigations show that these attain sexual maturity 



142 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 



WHALE 


LAM. 


NO. 


NO. 


675 


83 


1075 


85 


788 


76 


570 


76 


446 


75 


1120 


71 


1252 


70 


702 


67 


527 


66 


1356 


63 


1705 


62 


1855 


61 


1673 


60 


462 


60 


396 


59 


1744 


58 


1741 


58 


406 


56 


1627 


59 


1636 


56 


442 


59 


605 


55 


1113 


54 


1077 


54 


568 


54 


710 


53 


1978 


52 


1776 


52 


1745 


50 


1452 


50 


1112 


50 


1825 


49 


I860 


46 


1258 


46 


716 


45 


1121 


44 


1198 


44 


1326 


43 


976 


43 


927 


43 


1365 


42 


7861 


40 


1671 


40 




40 


1626 


39 


361 


38 


1597 


37 


1240 


36 


1450 


36 


1893 


36 


1666 


35 


1423 


35 


2120 


32 


1207 


31 


2083 


31 


1815 


31 


1888 


30 


1740 


30 


772 


30 


704 


30 


1239 


29 


1965 


29 


Mil 


23 


1599 


28 


2051 


28 


945 


27 


1822 


26 


1743 


26 


1674 


26 


1251 


25 


1286 


25 


363 


25 


1274 


24 


1451 


24 


1071 


24 


480 


24 


1635 


24 


2049 


23 


1902 


23 


2087 


23 


1774 


23 


2132 


23 


1777 


23 


1325 


23 



UNFUSED THICK 
CARTILAGE 



UNFUSED FINE 
CARTILAGE 



FUSED JOINTS 
INVISIBLE 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 143 



1287 

939 

857 

1355 

490 

1363 

523 

1081 

902 

1032 

1038 

521 

658 

654 

869 

1637 

531 

476 

1327 

1453 

1667 

1629 

2053 

1783 

1116 

1202 

1886 

525 

394 

1824 

650 

904 

356 

1894 

1821 

1205 

1663 

2005 

1203 

1742 

1243 

856 

558 

1206 

708 

1194 

2045 

1862 

2057 

931 

1244 

1290 

974 

1000 

941 

1589 

564 

1638 

1207 

1772 

517 

1850 

941 

900 

1857 

560 

1702 

1781 

1115 

1186 

440 

400 

1 1 10 

1932 

1818 



1972 

925 

560 

2041 

1700 

2047 

942 

1278 



Fig. 7. State of fusion of the vertebral epiphyes of a sample of male antarctic Fin whales 
Balaenoptera physalus. Each horizontal line represents a vertebral column which can be 

considered to be divided thus t A T t P T { L t C | where AT = anterior thoracic, 

PT = posterior thoracic, L = lumbar, C = caudal, 



i 4 4 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

at about 63 ft." The length at physical maturity has been ascertained at approxi- 
mately 68 ft. by Mackintosh & Wheeler, Peters, Brinkmann, Nishiwaki & Hayashi. 
In Text-fig. 3 the growth curve of the male whales cuts the 63 ft. ordinate at the 
9th lamination, but the upper part of the curve is almost horizontal and covers 
a wide range of lamination numbers at the average maximum length of 67-31 ft. 
Referring to the chart (Text-fig. 7) it will be seen that all the vertebral epiphyses are 
fused after the 36th lamination but as only two individuals are represented in the 
sample corresponding to the four numbers preceding 36, the actual number formed at 
physical maturity may be as low as 31. If the rate of laminae formation is approxi- 
mately 2 per year the male whales would appear to become sexually mature earlier 
than the females but take longer to become physically mature. 

The most recently established data about the ages of sexual maturity are those 
of Nishiwaki (1952) who quotes 3! years for males and 4 years for females. The 
present investigation gives a figure of 4 J years for males and 5-6 years for females. 
The discrepancy in these two results could be accounted for if the two most distal 
ear plug laminae could be regarded as being present in the foetus, i.e., the remnant 
of the epitrichium and the vernix caseosa. Against this hypothesis is the evidence 
of the ear plug of a 50 ft. long male, collected at Steinsham in 1955. This ear plug 
had only two laminae, one of which was the " foetal cone " Purves (loc. cit.) and 
was estimated by Laws & Purves (loc. cit.) to be one year old, and by Ruud, from 
inspection of the baleen plates to be 1 \ years old. The animal was of course much 
too long to be newly born. According to Sharpey-Schafer (1929) the vesicular 
cells of the epitrichium are shed late in the intra-uterine life and mingle with the 
secretion of the sebaceous glands to form a waxy covering, the vernix caseosa. The 
foetal cone of the ear plug is always very small, undifferentiated, mis-shapen, waxy 
and in the general appearance could represent the vernix caseosa. The penultimate 
lamination on the other hand, is always the deepest in the whole series and much 
too coarse to be present in the foetus. On the basis of the above estimation it 
represents from 1-1 J years' desquamation during the suckling and first free feeding 
life of the calf. The discrepancy could also be accounted for if the whales outside 
the " Sanctuary " become sexually mature earlier than those in Area I. 

It may be argued that the ovaries and baleen plates constitute an uncertain guide 
to the rate of formation of the laminae of the ear plug, so some alternative rates of 
formation must now be considered. In the first instance it must be stated that the 
extremely regular appearance of the laminae and the smooth, typically mammalian 
exponential growth curve afforded by plotting length against lamination number 
in both sexes of both samples preclude any possibility of a grossly irregular rate of 
desquamation. It is difficult to conceive of any cyclic phenomenon in the biology 
of whales or any other mammal for that matter which could account for three moults 
or desquamations per year. Whales like most other mammals are subject to a 
bi-annual rhythm of seasonal change and all the accumulated observations on their 
natural history suggest that their biology and physiology are adjusted accordingly. 
Any idea of an average rate of three desquamations per year derived from an irregular 
desquamation rate is precluded by the extremely regular appearance of the laminae. 
If four or more laminations per year are considered than the evidence of the ear 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 145 

plugs collected at Steinsham have some bearing on the problem. In that collection 
there was one plug (above cited) with only two laminations, one of which was the 
foetal cone. If the rate of formation of laminae had been four per year then that 
animal had grown from 21 ft. to 50 ft. in six months, and yet should theoretically 
have just been weaned at the time of capture. Now the length at weaning, of the 
Antarctic Fin Whale has been established by Mackintosh & Wheeler (1929) at 39 
ft. and since the North Atlantic Fin Whale is known to be smaller than the Antarctic 
form the average length at weaning of the former is likely to be less than 39 ft. 
A similar argument could be used for four of the Blue Whales represented in the 
present collection of ear plugs, since they possessed only two laminations and yet 
all the animals were over 70 ft. in length, i.e. some 20 ft. above the established length 
at weaning of the Blue Whale. 

Any rate of formation of laminae greater than four per year can be discounted on 
the basis of the above argument and there is left for consideration the idea of the 
formation of a single lamination per year. 

Assuming the formation of one ear plug lamination per year in the present sample, 
the ages at sexual and physical maturity of the female Fin Whales would be 10-12 
years and 30 years respectively and the oldest specimen some 85 years old. These 
figures are quite inconsistent with any of the previously assessed ages and growth 
rates of Fin Whales but are comparable with those in Man and require some critical 
examination. In the first instance it may be stated that the broad distal lamination 
was estimated to represent the combined weaning and first free feeding period of the 
animal's life, i.e. a period of approximately one year. The end of this period would 
correspond with the time of maximum blubber thickness for that year and would 
be followed by the northerly migration. If the subsequent desquamation of the 
whole body followed fluctuations in blubber thickness from year to year, the 
laminations of the ear plug would be formed annually. If, as seems likely, the ear 
plug is concerned in the conduction of sound from the blind end of the external 
auditory meatus to the middle ear, a desquamation rate correlated with blubber 
thickness would seem to be a necessity 

Reverting once more to the subject of ovaries — all authors are agreed that the 
onset of physical maturity corresponds with accumulation of 15-16 corpora lutea 
albicatia. If Rund's figure of i-6 corpora lutea per two years be used as the accu- 
mulation rate instead of that of Laws then the period between sexual and physical 
maturity would be 20 years and not 10 as previously estimated. In the absence of 
any definite proof of the rate of formation of the ear plug laminae, it can only be 
stated that the evidence shows conclusively that the rate is either one or two per 
year and that the latter is more consistent with all the previously accumulated 
knowledge of the natural history of Fin whales. 

THE AGE DISTRIBUTION 

Having accepted provisionally the validity of the hypothesis that the rate of 
increase in lamination-number is two per year, it is possible to form an age-frequency 
sample distribution from the sample frequencies of the lmination numbers. If 
" c" represents lamination number and, if it is assumed that the 21 th lamination is 



146 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

first recognizable approximately on the i th birthday, e.g. the 14th lamination on 
the 7th birthday, and the {21— i) th lamination approximately at age (i— J), then it 
is seen that whales with 2% laminations are between the ages of i and i + \ years 
and whales with (2i—i) laminations between the ages of (i— J) and i years. The 
sum of the two frequencies of lamination- numbers (2* — 1) and 2% is thus the sample 



20 
10 


FEMALES 




























* ^\. * * 
















N. • 
















• >w • • 
















^^. • 
















• ^Vs^^^ • 
















^ — _* * * 

















• •** -_ 




• 


• 


* 






MALES 
















* 














to 


\ * • 


• 















• - 




L t 


• . 


. 


• 










10 Z ° VEAtS 


30 






40 







Fig. 8. Smoothed age frequency distribution of a sample of fin 
whales Balaenoptera physalus 



frequency of the one year group of individuals whose nearest birthday is * years. 
In the case of the females sampled in Area I only the frequencies of lamination- 
numbers of 9 or more were used in the construction of the age-frequency distribution, 
it was considered, because of the corresponding smaller average total body lengths, 
that the smaller lamination numbers were not fully represented in the sample. The 
age-frequencies of the Area I females are shown in Text-fig. 8. 

At the outset it must be emphasized that owing to the wide variability of the data, 
the small size of the sample and the possible lack of randomness of selection, due 
to differences in capture proneness, etc., no more is claimed for the ensuing analysis 
than that any conclusions based on it are but crude approximations of the true values 
of the population. 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 147 

The simplest hypothesis to fit the data is that the mortality rate is constant, i.e. 
the probability that an individual will survive a further year is independent of its 
present age. In this case the age frequencies follow a geometrical progression and the 
points on the graph of the frequency distribution lie on a curve of exponential form. 
Fitting by weighted least squares, the curve : 

a, _ gZ .45-0. 133 x 

was obtained, where % is the age in years and is seen in Text-fig. 8 to give a fairly 
good description of the sample frequencies. This curve implies that over the given 
range of age 5 or more years a proportion £-°- 133 or approximately 88% of each year 
group survives to the following year. The insensitive nature of this estimate of 
survival rate is demonstrated by its 95% confidence interval which ranges from 

79% to 97%. 

In order to discount the under-representation of the smaller whales the above 
survival curve was constructed using only the data for not less than 9 laminations 
at which according to the growth curve in Text-fig. 3 the mean length is 65 ft. The 
same minimum average length is insured in the sample of males captured in Area 
I by using only the data of 13 or more laminations. The age-frequencies for the 
Area I males are shown in Text-fig. 8. As in the case of the females a weighted 
least squares exponential curve was fitted to the data and as is seen in Text-fig. 8 
the curve : 

V = ^3 .23-0. 106 x 

does reflect the characteristics of the sample frequencies. This curve implies a 
survival rate of <?-°- 106 or 90% for each year group. There is apparently no evidence 
of a significant difference in survival rates between the two sexes. 

Assuming the perhaps somewhat unreal hypothesis of a stable population, constant 
in size, it is possible to conjecture estimates of the immature females' mortality 
rate. The estimation of this mortality rate will be made for each of the following 
four models of birth rate. 

Model 1. First offspring at approximately 5 years old with a succeeding 
birth rate of one offspring produced every 2 years. 

Model 2. First offspring at 6 years old with succeeding birth rate of one 
offspring every two years. 

Model 3. First offspring at 5 years old with succeeding birth rate of one 
offspring every three years. 

Model 4. First offspring at 6 years old with succeeding birth rate of one 
offspring every three years. 

Under the conditions of Model I the expected number of female offspring, assuming 
that the birthrate is the same for the two sexes, is at the rate of one every 4 years. 
The total number of female offspring per year is then expected to be approximately : 

2 Bl(i) x J 
• *- 5 

where Bl(i) equals the number of females in the age group (i — -|) to (i + \) years 

old. If B is assumed to be the constant yearly number of female births, then l(i) 



148 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

is seen to be the approximate probability that a female will survive to an age of i 
years. Equating the two expressions of total yearly female births 

S Bl{i) xi = B 

*= 5 

it follows that 

S l(i) = 4. 
*= 5 

Now the survival probabilities l(i) are estimated as being proportional to the yearly 

ordinates of the fitted age distribution curve y = ^ .45-0.133 x 

i.e. 



where 



'« = 4^ 



yj __ g3.45-0.133i 



(*' > 5) 



The survival probabilities for the other 3 models may similarly be obtained and 
the numerical values of the l(i) for all 4 models are tabulated in Table I. Thus, 
for example, it is seen from this table that under the conditions of Model 1 there is 

Table I. — Provisional Mortality Rates of Antarctic Fin Whales 









Model 1 


Model 2 


Model 3 


Model 4 




Observed 
3 i frequencies 


Fitted 
frequencies 


a 










..-^,. 




Ag 


100 l(i) 100 d{i) 


100 l(i) 


100 d(i) 


100 l(i) 


100 d(i) 


t \ 
100 l(i) 100 d(i) 


Undei 


5 • — 


— 


— 50-0 


— 


— 


— 


25-0 


— 


— 


Undei 


•6 . — 


— 


— — 


— 


50-0 


— 


— 


— 


25-0 


5 


20 


. 16-130 


50-0 6-2 


— 


— 


75-o 


9-4 


— 


— 


6 


14 


14-116 


43-8 5'5 


50-0 


6-2 


65-6 


8-2 


75 -o 


9'4 


7 


10 


• 12-353 


38-3 4-8 


43-8 


5-5 


57-4 


7 -i 


65-6 


8-2 


8 


11 


. 16-810 


33*5 4*2 


38-3 


4-8 


50-3 


6-3 


57-4 


7-1 


9 


7 


9-460 


29-3 3-6 


33*5 


4-2 


44-0 


5-5 


50-3 


6-3 


10 


6 


8-279 


25.7 3.2 


29*3 


3-6 


38-5 


4.8 


44-0 


5-5 


11 


5 


7*245 


22-5 2-8 


25-7 


3*2 


33*7 


4-2 


38-5 


4-8 


12 


8 


6-340 


19.7 2-5 


22-5 


2-8 


29*5 


3'7 


33*7 


4-2 


13 


7 


- 5-548 


I7-2 2-1 


19-7 


2'5 


25-8 


3-2 


29*5 


3-7 


14 


6 


- 4-855 


15-1 1-9 


17-2 


2-1 


22-6 


2-8 


25-8 


3'2 


15 


3 


4-249 


13-2 1-7 


i5-i 


1-9 


19-8 


2-5 


22-6 


2-8 


16 


5 


- 4-7i8 


n-5 1-4 


13-2 


1-7 


17-3 


2-1 


19-8 


2'5 


17 


3 


• 3-254 


io-i 1-3 


n-5 


1-4 


i5-i 


1-9 


17-3 


2-1 


18 


7 


2-848 


8-8 i-i 


IO-I 


i-3 


13*2 


1-6 


i5-i 


i-9 


19 


1 


2-492 


y-7 0-9 


8-8 


1 -i 


n-6 


i-5 


13-2 


i-6 


20 


6 


2-181 


6-8 o-8 


7-7 


0-9 


IO-I 


i-4 


n-6 


i-5 


21-25 


7 


• 7-438 


4-6 3-6 


5*3 


3-9 


6-9 


5-2 


8-o 


6-o 


26-30 


6 


. 3-8i6 


2-4 1.5 


2-7 


1-7 


3-5 


4 -i 


4-i 


2-5 


30-40 


3 


2-966 


0-9 0-9 


1 -o 


1 -o 


1-4 


i-6 


i-6 


1-6 




Note. The above mortality rates 


> were calculated 


from the 


age freqi 


lency distribution 


of a singl 


e 



sample of the population using the ear plug lamination as guide to the age. The figures are based 
on an assumed rate of formation of 2 laminations per year. If the rate of formation is one lamination 
per year the age increments in column I would have to be doubled. 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 149 

an estimated proportion of 50% surviving to an age of 5 years and a proportion of 
257% surviving to an age of 10 years. The value d(i) given in the same table are 
calculated from the relationship 

d(i) = l(i) - l(i + 1) 

and therefore 100 d(i) is the estimated percentage of the population which dies 
between the ages of i and (i + 1) years. Thus in Model 1 an estimated 5*5% 
die between the age of 6 and 7 years. From the columns of d(i) it is seen that 
the immature mortality rates are such that 50% of the population dies under the age 
of 5 in Model 1, 50% die under the age of 6 in Model 2, 25% die under the age of 
5 in Model 3, and in Model 4, 25% die under the age of 6. It is of some interest to 
note that if the fitted exponential curve is extra polated backwards to age zero then 
the ensuing immature death rate is such that 487% of the population die under 
the age of 5 years, and 55% under the age of 6 years, values very close to that obtained 
under Models 1 and 2. 

From the growth curve of Area I females (Text-fig. 3) it is seen that the fitted 
regression function estimates the mean total lengths at ages 3, 4 and 5 years as 
61-5 ft., 64-0 ft. and 66-o ft. respectively. As the whales prone to capture are restricted 
to a minimum length of 57 ft., there is a strong likelihood that the given mean values 
of total length of the 3- and 4-year-olds are over-estimates of the population values 
at these ages, but as the range of the sample of the 5-year-olds (9 and 10 laminations) 
is above 6o-o ft. there is no reason to believe that the estimation of this age group 
has been affected by the size restriction. If the population proportional frequencies 
of the 3 and 4 years old are assumed to be given by the extrapolated values of the 
exponential female age-distribution (Text-fig. 8) then as 5 3-year-olds and 14 4-years 
are present in the sample it may be inferred that approximately 23% of the 3-year- 
olds and 76% of the 4-year-olds are over 57 ft. in length, and thus prone to capture. 
As stated before, the sample evidence indicates almost conclusively that 100% of 
the 5-year-old age group are over 57 ft. 

The figure for the percentage of immature whales in the total catch is, of course, 
dependent upon the age of attaining sexual maturity ; this age is assumed to be 
4 years in Models 1 and 3, and 5 years in Models 2 and 4. The numbers of females 
taken in Area I under the ages of 5 and 6 years are respectively 39 and 19 out of a 
total catch of 156 whales giving the percentage of immatures in the catch as 12-2 
for Models 1 and 3 and 25-0 for Models 2 and 4. 

CONCLUSIONS 

The original hypothesis of a bi-annual rhythm for the formation of the laminae 
having been supported by correlation with observations on the growth of the baleen 
plates up to the sixth year of life it remained to establish whether this rhythm 
continued throughout the life of the animal. 

It has been shown that the growth of the plug is not directly associated with the 
lateral growth of the skull, but that there is an exponential relationship between 
the total body length and the lamination number. 

The exponential growth curve of the body length approximates in form to the 



i 5 o EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 




S1V/IOIMQNI 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 151 

normal mammalian growth curve and it was therefore assumed that the laminae 
of the ear plug were formed at regular intervals throughout life. 

Assuming the interval between sexual and physical maturity to be 10 years as 
assessed from previously established information about the number and rate of 
formation of corpora lutea produced during this interval, it has been shown that the 
rate of formation of laminae is approximately 2 per year. 

The previously established total body lengths of female Fin Whales at sexual 
and physical maturity, i.e., 66 ft. and 72*4 ft. respectively are identical with those 
deduced from the ear plug body length growth curve using the above rate of laminra 
formation. 

The fact that the average core length per lamination number is almost identical 
in the two sexes, notwithstanding the difference in the skull proportions, indicates 
that the method of counting is fairly accurate. 

Using the lamination number as a guide to age, the age frequency distribution 
of the sample takes the form of exponential curves in both sexes. From these 
exponential curves an age independent adult mortality rate of 12% per annum 
has been deduced for the female population, and 10% for the males. 

The age dependent mortality rates for the female population have been tabulated 
using two different ages at sexual maturity and two durations of breeding cycle. 
Assuming the age at sexual maturity to be five years with the first offspring at six 
years the immature female mortality rate would be approximately 50%. 

Assuming the above age at sexual maturity and that the Area I sample is repre- 
sentative of the catch then 25% of the catch is composed of immature specimens ; 
a figure which accords well with the average annual catch of immature animals and 
the expected proportion prone to capture under the existing regulations. It is 
interesting to note that the female adult mortality rate of S. Georgia population 
between the years 1925-31 was assessed by Wheeler (1934) at 13% per annum and 
the theoretical immature mortality rate at approximately 50% ; it will be seen that 
the adult female mortality rate for the present Area I sample was found to be in 
close agreement with these figures. With the age at sexual maturity at 5 years and 
the fertility rate 25% the above figures are exactly those required to keep the popu- 
lation stable in size. They also approximate to the figures for the apparent mortality 
rate which would be obtained by sampling an increasing population with a negligible 
mortality rate. 

From the statistical analyses made by Hylen, Jonsgard, Pike & Ruud (1955) 
it may be noted that the peak catches are obtained according to the baleen plate 
data from age group 3 for females and age group 4 for males. According to the 
ear plug data from Area I the peak catches were obtained from the age groups 4-6 
years in respect of females and 6-8 years in respect of males. This discrepancy 
may be partly explained by a difference between the mean total body lengths in 
the two populations, but is probably mainly due to the increased proportion of young 
animals in the population outside the " Sanctuary " the adult mortality rate of which 
was estimated to be 25%. 

From the Hylen et al. analysis it appears that the population has become sexually 
mature at an earlier age in recent years. If Jonsgard's (1952) body length figure 



152 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

of 67 ft. at sexual maturity is applied to the growth curve of the Area II sample 
it will be seen to cut the x ordinate at the 8th lamination giving an age at sexual 
maturity of 4 years. 

The oldest specimen recorded in the present sample was 42 years old and in this 
respect it is necessary to draw attention to a paper by Simons (1957) in which he 
states that in an unmolested Humpback Whale population the life span was very 
low. He bases his comments on the fact that only a small proportion of his sample 
of 146 individuals had attained an age greater than 14 years and that only 1 had 
reached the age of 29. It will be noted that his sample size for both sexes is smaller 
than the Area I sample. Under the conditions of Model 1 only 13% of the Area I popu- 
lation is over 14 years old and only 2-4% over 25 years old ; the latter figure embraces 
all ages between 26 and 30, so the figure for the 29-year-olds is more likely to be 
J%. These figures give no exact indication of the maximum life span of the animals. 
The question as to whether or not the material described in the present paper repre- 
sents a random sample can be judged from Text-figs. 9 & 10. In Text-fig. 9 the 
age frequencies are plotted at intervals of two years, i.e. the approximate breeding 
cycle. It will be seen that males and females are present in each group in comparable 
numbers except that the peak value for males occurs two years later than that of the 
females. The left-hand side of the curve is much steeper than that of the right in 
both sexes notwithstanding that all the animals represented are above the permitted 
size limit. This effect is not due to any lack of randomness in the sample, nor to 
any length discrimination on the part of the gunners in the whaling fleet, both of 
which ideas have been suggested to explain the phenomenon in other samples. 
The steepness of the left-hand side of the curve is due to the fact that in the immature 
age groups the length variation is both relatively and absolutely greater than that 
of the adults, the standard deviation per age group in the sample being in the region 
of 5 ft.; thus a significant proportion of the immature animals is below the permitted 
size limit and therefore absent from the sample. As might be expected this effect 
is observable at a greater age in the males than in the females. In Text-fig. 10 
the total frequencies of male and females have been plotted at two-yearly intervals. 
The dotted line represents the age frequency distribution of a hypothetical population 
of whales breeding every two years in which every individual becomes sexually 
mature at six years and in which the age frequency distribution remains stable with 
a constant mortality rate. Such a hypothetical population is perhaps unrealistic 
but the shape of the curve does indicate that the age frequency distribution of the 
sample approximates to that of a theoretically possible whale population. It is 
reasonable to assume that the sample is randomly drawn from a population with 
the above biological characteristics. If Simon's material is assumed to represent 
a truly random sample of the population then it would be statistically unlikely 
for more than one specimen in the sample of 146 females to be 29 years old, even if 
the maximum life span was 42 years or more. 

Of the 12 Blue Whale ear plugs in the present collection only two were taken 
from animals more than 2 years old ; 1 was from an animal 6 years old and the 
other 12 years old. 

The above figures are based on the assumption that the ear plug laminae are 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 153 

formed at the rate of 2 per year but with the possible formation of one lamination 
per year the values would have to be doubled. For the purpose of estimating changes 
in the population structure the actual rate of laminar formation would appear to 
be immaterial provided the error, if any, is constant from year to year and age 
group to age group. From this point of view the ear plug is probably a more reliable 



80 
78 


, 






76 


\ 






74 


\ 






72 


\ 






70 


\ 






68 


\ 






66 


\ 






64 


\ 






62 


\ 






60 


\ 






58 


\ 






"J 66 


\ 






- S4 


J \ \ 






< 52 


/■ 1 1 






£ so 


/ \ \ 






U. 48 


/ v \ 






A <* 


/ • \ 






2 44 


/ M 






< 42 








<n 40 








3 38 


/ r 






< 36 


/ \\ 






Z 34 








a 32 








30 


1 \ /*> \ 






</) 28 


/ ^ ^ \ 






i* 


x \ 






»- 24 


\\ 






O 22 


\\ 






20 
18 


x \ 






16 


\X 






14 


\ \ 






12 


V, 






10 




"*-v\ / 




8 








6 








4 




^**-->- 




2 






s^.^^ ^*^"" ■* - ^ ^^" ■■ 



10 II 14 16 



18 20 22 24 26 28 30 32 34 36 38 40 42 
AGE, IN YEARS 



Fig. 10. Age distribution of a sample of fin whales Balaenoptera physalus. 
Sample distribution = X. Theoretical population distribution = o. 



guide to age than the ovary. The assumption on which the life tables were estimated 
i.e. that the size of the population is a constant number from year to year, is clearly 
to be regarded as no more than a crude first approximation to the actual form of 
population growth. However, with the present set of data it is necessary, for 
estimation purposes to make some such an assumption for it is not possible to deduce 
from data of one single year whether the population size is increasing, decreasing 
or static. It is evident that a more refined analysis of population growth can be 
applied only to extensive acts of data obtained in several successive years. 



154 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

ACKNOWLEDGMENTS 

Grateful acknowledgments are due to H.M. Ministry of Agriculture and Fisheries 
Whaling Inspectors aboard the factory ships Balaena, Southern Harvester and Southern 
Venturer during the Pelagic Whaling Season 1955-56 upon whose collections this 
report is based. Thanks are due to the National Institute of Oceanography for 
making available to us the material so collected. We wish particularly to express 
our appreciation to Mr. M. R. Clarke for the excellent set of data provided in his 
log, of which only the most obviously relevant details have been reproduced in our 
tables. We should also like to thank Messrs D. W. Cooper and M. G. Sawyers for 
the microscope and photomicrographic work involved and Miss J. R. Proctor for 
carrying out most of the computation. Finally we wish to thank Mr. J. G. Skellem 
and Dr. F. C. Fraser for their helpful criticism. 

Masaharu Nishiwakis' " Age characteristics of Ear Plugs of Whales " reprinted 
from the Scientific Reports of the Whales Research Institute, No. 12, 1957 was received 
after completion of the foregoing account. It is gratifying to see that his results, 
based on a much smaller sample are in general agreement with the above. His 
new figure of 64 ft., for the age at sexual maturity of female Fin Whales does not 
coincide with previously published figures but if applied to Text-Fig. 3 of the present 
paper, it would give an age at sexual maturity of 4-5 years, which is identical with 
his own published result. 



REFERENCES 

Brinkmann, A. 1948. Studies on female Fin and Blue Whales. Hvalradets Skrifter, 31 : 

1-38, 13 figs. 
Brown, S. G. 1954. Dispersal in Blue and Fin Whales. " Discovery " Reports, 26 : 335-384, 

14 figs. 
Hylen, A., Jonsgard, A., Pike, G. C. & Ruud, J. T. 1955. The age composition of Antarctic 

Fin Whale catches. The Norwegian Whaling Gazette, 10 : 577-589, 2 figs. 
Jonsgard, A. 1952. On the growth of the Fin Whale in different waters. Ibid., 41 : 58-65, 

5 fi gs. 
Laurie, A. H. 1937. The age of female Blue Whales. " Discovery " Reports, 15 : 223-284, 

14 figs. 
Laws, R. M. 1956. Breeding and mortality rates of Antarctic Fin Whales. Abstracts of the 

Challenger Society, 3 : No. 8. 
Laws, R. M. & Purves, P. E. 1956. The ear plug of the Mysticete as an indication of age, 

with special reference to the North Atlantic Fin Whale. The Norwegian Whaling Gazette, 

45 : 413-425. 12 figs. 
Mackintosh, N. A. & Wheeler, J. F. G. 1929. Southern Blue and Fin Whales. " Discovery" 

Reports. 1 : 257-540, 21 pis., 157 figs. 
Mackintosh, N. A. 1942. The southern stocks of Whalebone Whales. Ibid., 22 : 197-300, 

9 figs. 
Nishiwaki, M. & Hayashy, K. 1950. A biological survey of Fin and Blue Whales. The 

Scientific Reports of the Whales Research Institute, 4 : 132-190, 57 figs. 
Nishiwaki, M. & Oye T. 195 1. A biological investigation on Blue and Fin Whales caught by 

Japanese Antarctic Whaling Fleet. Ibid., 5 : 91-169, 36 figs. 
Peters, N. 1939. Uber Grosse, Wachsturn und Alter des Blauwales und Finnwales. Zoo- 

logischer Anzeiger, Bd., 127 : 193-204, 3 figs. 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 155 

Purves, P. E. 1955. The wax plug in the external auditory meatus of the Mysticeti. 

" Discovery " Reports, 27 : 293-302, 4 pis., 2 figs. 
Simons, H. W. 1956. Some observations on the ear of Blue and Fin Whales. The Norwegian 

Whaling Gazette, 45 : 37-42, 2 figs. 
Simons, H. W. & Weston, R. D. 1957. An underfinished Humpback population ? Ibid., 

46 : 231-238. 
Schafer, E. S. 1929. The Essentials of Histology, 12th Edition London, 1929. Longman and 

Green. 
Ruud, J. T. 1949. Further studies on the structure of the baleen plates and their application 

to age determination. Hvalradets Skrifter, 29 : 1-69, 28 figs. 
Van Oosten, J. 1957. The skin and scales. The Physiology of Fishes. 1 : 207-243, 10 figs. 

Edited by M. E. Brown. 
Wheeler, J. F. G. 1934. The stock of whales at South Georgia. " Discovery " Reports, 

9 : 351-372, 3 figs. 

APPENDIX 
Table A. — Female Antarctic Fin Whales Balaenoptera physalus Sample Area I 









Esti- 


Length 








Esti- 


Length 




Total 


Number 


mated 


of 




Total 


Number 


mated 


of 


Whale 


length 


of 


age 


core 


Whale 


length 


of 


age 


core 


number 


(ft.) 


laminae 


(years) 


(mm.) 


number 


(ft.) 


laminae 


(years) 


(mm.) 


1284 


58 


5 


■ 21 


■ 58 


519 


. 60 


10 


5 


• 34 


863 


63 


6 


■ 3 


■ 32 


2043 


. 60 


10 


• 5 


• 23 


1714 


65 


6 


• 3 


. 27 


1749 


. 63 


10 


5 


• 37 


352 


67 


6 


3 


■ 36 


1898 


■ 63 


10 


5 


35 


1820 


68 


6 


3 


25 


1247 


. 66 


10 


5 


• 30 


2013 


58 


7 


■ 34 


■ 58 


859 


• 67 


10 


• 5 


• 35 


1200 


59 


7 


34 


■ 59 


1279 


. 69 


10 


• 5 


• 34 


1850 


61 


• 7 


■ 34 


• 30 


1196 


■ 73 


10 


5 


■ 33 


739 


63 


7 


34 


■ 30 


1937 


. 62 


11 


54 


. 14 


1785 


64 


7 


34 


■ 25 


515 


. 64 


11 


54 


— 


1819 


68 


74 


31 


35 


1890 


65 


11 


54 


40 


648 


69 


7 


34 


37 


1821 


65 


11 


54 


62 


601 


69 


7 


34 


45 


1817 


66 


10 


5 


33 


1707 


58 


8 


4 


58 


935 


66 


11 


54 


30 


652 


58 


8 


4 


58 


1 165 


68 


11 


54 


46 


776 


60 


8 


4 


25 


1167 


61 


12 


6 


18 


1856 


60 


8 


4 


30 


1208 


63 


12 


6 


34 


871 


65 


8 


4 


30 


448 


67 


12 


6 


44 


1454 


66 


8 


4 


26 


1320 


67 


12 


6 


44 


2059 


60 


9 


44 


— 


io34 


67 


12 


6 


35 


1864 


60 


9 


44 


33 


1209 


68 


12 


6 


49 


1930 


61 


9 


44 


3i 


786 


73 


12 


6 


45 


2126 


62 


9 


44 


27 


2095 


76 


12 


6 


40 


1748 


63 


9 


44 


24 


1812 


64 


13 


64 


— 


1108 


64 


9 


44 


48 


2097 


65 


13 


64 


— 


933 


65 


9 


44 


33 


670 


72 • 


13 


6* . 


50 


1118 


65 


9 


44 


— 


656 


67 


14 


7 


36 


1248 


67 


9 


44 


37 


2128 


67 


14 . 


7 


40 


1036 


68 


9 


44 


40 


1968 


68 


14 . 


7 ■ 


40 


1813 


68 


9 


44 


33 


1784 


68 


14 


7 - 


35 



156 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

Table A — cont. 









Esti- 


Length 








Esti- 


Length 




Total 


Number 


mated 


of 




Total 


Number 


mated 


of 


Whale 


length 


of 


age 


core 


Whale 


length 


of 


age 


core 


number 


(ft.) 


laminae 


(years) 


(mm.) 


number 


(ft.) 


laminae 


(years) 


(mm.) 


906 


70 


14 


7 


34 


529 


70 


26 


• 13 


• 53 


1901 


70 


14 


7 


34 


1625 


70 


. 27 


• I3i 


. 42 


1079 


72 


14 


7 


— 


1816 


73 


27 


• 13* 


■ 73 


1811 


67 


15 


74 


40 


1704 


73 


27 


• 13* 


• 9i 


660 


7i 


15 


74 


52 


1040 


68 


28 


■ 14 


■ 78 


855 


72 


15 


74 


43 


743 


74 


28 


• 14 


■ 78 


1285 


70 


16 


8 


53 


1 169 


68 


29 


. 14* 


94 


1775 


7i 


16 


8 


34 


747 


69 


29 


14* 


42 


2044 


72 


16 


8 


50 


1966 


73 


30 


15 


70 


7°3 


72 


16 


8 


60 


1280 


70 


3i 


154 


75 


1675 


72 


16 


8 


40 


672 


70 


32 


16 


80 


1927 


72 


16 


8 


40 


1710 


7i 


32 


16 


66 


562 


73 


16 


8 


56 


1630 


74 


32 


16 


87 


1823 


77 


16 


8 


42 


486 


75 


32 


16 


52 


513 


69 


17 


i8£ 


37 


2134 


76 


32 


16 


55 


737 


7i 


17 


H 


60 


1628 


75 


33 


i6£ 


60 


1854 


74 


17 


8} 


32 


1633 


67 


34 


17 


74 


784 


68 


18 


9 


25 


482 


75 


34 


17 


70 


896 


7i 


18 


9 


34 


780 


75 


35 


174 


63 


2136 


7i 


18 


9 


42 


74i 


67 


36 


18 


70 


1085 


73 


18 


9 


49 


712 


72 


36 


18 


67 


1964 


74 


19 


9i 


47 


1933 


72 


36 


18 


45 


898 


70 


20 


10 


50 


2007 


72 


36 


18 


65 


597 


70 


20 


10 


40 


593 


73 


36 


18 


63 


2099 


70 


20 


10 


50 


1669 


75 


36 


18 


33 


674 


72 


20 


10 


5i 


i93i 


76 


37 


18* 


50 


1664 


72 


20 


10 


60 


1670 


73 


39 


194 


73 


646 


74 


21 


10* 


63 


1044 


68 


40 


20 


60 


1083 


70 


22 


11 


34 


2015 


70 


40 


20 


75 


1779 


7i 


22 


11 


38 


937 


72 


40 


20 


60 


2122 


72 


22 


11 


29 


1706 


72 


40 


20 


94 


714 


7i 


22 


11 


70 


488 


73 


40 


20 


90 


1739 


73 


23 


Hi 


72 


2085 


72 


42 


21 


70 


894 


7i 


24 


12 


56 


1042 


74 


42 


21 


92 


1788 


7i 


24 


12 


— 


2130 


69 


43 


2i4 


53 


1892 


72 


24 


12 


— 


438 


72 . 


45 


22£ 


58 


700 


72 


24 


12 


27 


906 . 


75 ■ 


46 


23 


145 


603 


73 


24 


12 


53 


1634 


72 


50 


25 


73 


1366 


73 


24 


12 


— 


1425 


81 


50 


25 


39 


434 


78 


34 


12 


80 


1250 


69 


54 


27 


75 


398 


67 


24 


12 


67 


1357 


77 


54 


27 


96 


595 


7i 


25 


12* 


— 


1703 


69 


55 


27i 


69 


749 


7i 


25 


12} 


59 


1 163 


70 


56 


28 


60 


745 


70 


26 


13 


— 


1938 


72 


56 


28 


7i 


2099 


70 


26 


13 


50 


1352 


72 


56 


28 


56 


867 


72 


26 


13 


17 


2009 


75 


66 


33 


93 


1358 


74 


26 


13 


74 


706 


75 


70 


35 


88 


478 


77 


26 


13 


85 


358 


73 


80 


40 . 


120 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 157 



Table B. — Male Antarctic Fin Whales Balaenoptera physalus Sample Area I 



Total 

length 

(ft.) 



Number 

of 
laminae 



Esti- 
mated 
age 

(years) 



• 55 


3* 


T 3 

I4 


• 58 


4 


2 


. 60 


4 


2 


. 64 


6 


3 


■ 58 


7 


3* • 


• 58 


7 


3* • 


. 68 


8 


4 


. 61 


8 


4 


. 61 


9 


4* • 


. 61 


9 


4i • 


. 62 


9 


4* • 


• 63 


9 


4* • 


. 64 


9 


4* • 


. 64 


9 


4i • 


• 67 


9 


4* • 


• 65 


9 


4* ■ 


. 61 


10 


5 


• 63 


10 


5 


. 66 


10 


5 


. 66 


10 


5 


. 67 


10 


5 


. 62 


11 


5i • 


• 63 


11 


5* ■ 


. 69 


11 


5i • 


• 63 


12 


6 


. 64 


12 


6 


• 65 


12 


6 


. 66 


12 


6 


. 68 


12 


6 


. 62 


13 


6* . 


. 64 


13 


6* • 


. 64 


13 


6* . 


• 65 


13 


6* . 


• °5 


13 


6* • 


• 65 


13 


6* . 


• 67 


13 


6* . 


• 69 


13 


6* . 


. 62 


14 


7 


. 62 


14 


7 


• 65 


14 


7 


• 65 


14 


7 


. 66 


14 


7 


. 66 


14 


7 


• 67 


14 


7 


. 67 


14 


7 


. 68 


14 


7 


. 64 


15 


7* • 



Length 

of 

core 

(mm.) 



20 

26 

14 

27 

58 
24 

30 
4i 
3 1 
4 1 
24 
38 
34 
25 
3i 
56 
25 
68 

32 
43 
43 
29 

4i 
38 
30 
45 
30 
60 

47 
40 

37 
42 
37 
43 
38 
43 
48 

48 
35 
27 
55 
30 
13 
42 
38 



Total Number 
Whale length of 

number 
1827 
1894 

356 

904 

650 
1824 

394 

525 
1886 
1202 
1116 
1783 
2053 
1629 
1667 

1453 
1327 

476 

531 
1637 

869 

654 

658 

521 
1038 
1032 

902 
1081 

523 
1363 

490 

1355 

857 

939 
1257 
1671 
1861 
1356 

927 

976 
1326 
1198 
1121 

716 
1238 
i860 
1825 



Esti- 
mated 
age 



(ft.) 


laminae 


(years) 


. 66 


15 


74 • 


. 66 


15 


7* • 


. 66 


15 


7i ■ 


. 67 


15 


7i • 


• 63 


16 


8 


■ 65 


16 


8 


• 6 5 


16 


8 


. 66 


16 


8 


. 67 


16 


8 


• 67 


16 


8 


. 67 


16 


8 


. 68 


16 


8 


. 68 


16 


8 


• 65 


17 


8* . 


. 66 


17 


8* . 


• 67 


17 


8* . 


. 67 


17 


8* • 


. 69 


17 


8* . 


• 67 


17 


8* • 


• 69 


17 


8* . 


. 69 


18 


9 


. 66 


19 


9i • 


• 7i 


19 


9* • 


• 65 


20 


10 


. 66 


20 


10 


. 66 


20 


10 


• 53 


20 


10 


• 67 


20 


10 


. 68 


21 


104 


• 73 


21 


io£ 


. 66 


22 


11 


. 66 


22 


11 


. 68 


22 


11 


• 7 1 


22 


11 


• 7 1 


22 


11 


. 67 


40 


20 


• 70 


40 


20 


. 66 


42 


21 


. 66 


43 


21* ■ 


. 68 


43 


2l£ 


• 65 


43 


214 . 


. 66 


44 


22 


. 66 


44 


22 


. 68 


45 


22^ 


. 66 


46 


23 


. 68 


46 


23 


• 67 


49 


24i • 



Length 
of 

core 
(mm.) 

42 
27 
43 
50 
36 
60 
70 
48 
53 
63 
35 
45 
55 
60 

55 
35 
45 
50 
50 
50 
69 
38 
57 
26 
62 
53 
53 
40 

40 

46 
62 
60 
54 

50 
100 

77 
78 
77 
42 
80 
50 
38 
85 
94 
65 



158 EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 

Table B — cont. 









Esti- 


Length 






Total 


Number 


mated 


of 




Whale 


length 


of 


age 


core 


Whale 


number 


(ft.) 


laminae 


(years) 


(mm.) 


number 


III2 


. 64 


• 5o 


• 25 


• 65 


1902 


1452 


. 68 


50 


• 25 


• 50 


2049 


1745 


• 71 


50 


• 25 


. s 5 


1635 


I776 


■ 65 


• 52 


. 26 


• 52 


480 


1978 


• 67 


52 


. 26 


. 68 


1071 


7IO 


■ 65 


53 


26£ 


65 


I45i 


568 


62 


54 


27 


88 


1274 


IO77 


66 


54 


27 


46 


363 


III3 


68 


54 


27 


58 


1286 


605 


66 


55 


2 7 £ 


83 


1251 


442 


7i 


59 


28£ 


63 


1674 


1636 


63 


56 


28 


62 


1743 


1627 


65 


59 


28£ 


90 


1822 


406 


65 


56 


28 


62 


945 


I74I 


65 


58 


29 


67 


2051 


1744 


68 


5^ 


29 


40 


1599 


396 


67 


59 


29i 


77 


IIII 


662 


66 


60 


30 


85 


1965 


1073 


72 


60 


30 


120 


1239 


1855 


68 . 


61 


3°h - 


65 


704 


i7°5 • 


73 • 


62 


3i 


80 


772 


1353 


70 


63 


31* 


50 


1740 


527 


65 • 


66 


33 


80 


1888 


702 


70 . 


67 . 


33i 


68 


1815 


1252 . 


73 ■ 


70 


35 


— 


2083 


1120 . 


65 • 


71 • 


35i 


100 


1207 


416 


68 . 


75 • 


37h • 


— 


2120 


57° ■ 


67 . 


76 . 


38 


130 


1423 


788 . 


66 . 


76 . 


38 


84 


1666 


1075 • 


64 . 


85 • 


4 2 i • 


72 


1893 


676 . 


64 . 


83 • 


41* ■ 


95 


1450 


1325 • 


64 . 


23 


n| . 


42 


1240 


1777 • 


65 • 


23 


ill . 


46 


1597 


2132 . 


65 • 


23 


uj . 


40 


361 


1774 • 


67 . 


23 


ii-i- . 


65 


1626 


2087 . 


68 . 


23 


ii* . 


33 


1671 



Total 

length 

(ft.) 

68 

71 
64 

67 
68 
70 
74 
63 
67 
70 
67 
65 
79 
64 
66 
63 
67 
69 
68 
70 

65 
66 
66 
69 
65 
68 

65 
7° 
66 

64 
69 
70 

7i 
65 
67 
67 



Number 

of 
laminae 

23 
23 
24 
24 
24 
24 
24 
25 
25 
25 
26 
26 
26 
27 
27 
28 
28 
28 
29 
29 
30 
30 
30 
30 
31 
3i 
32 
33 
35 
36 
36 
36 
36 
37 
38 
40 



Esti- 


Length 


mated 


of 


age 


core 


(years) 


(mm.) 


iii 


. — 


ui 


• 57 


12 


• 35 


12 


■ 58 


12 


• 55 


12 


. 40 


12 


• 45 


12I 


• 45 


12* 


• 43 


12* 


■ 56 


13 


43 


13 


— 


13 


45 


13* 


57 


13* 


53 


14 


63 


14 


62 


14 


62 


14* 


82 


Hi 


— 


15 


65 


15 


48 


15 


83 


15 


47 


I5i 


70 


15* • 


68 


16 


60 


16* . 


40 


17* • 


33 


18 


58 


18 


80 


18 


80 


18 


40 


i8i 


48 


19 


60 


20 


50 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 159 



Table C. — Female Antarctic Fin Whales Balaenoptera physalus Sample Area II 
Whale Total Number Estimated Length of 



number 


length (ft.) 


of laminae 


age (years) 


core (mm.) 


Factory ship 


1480 


60 


3 


1* 


20 


Southern Venturer 


A1493 


62 


5 


2* 


25 


> > t> 


A1231 


67 


5 


2* 


30 


Balaena 


— 


63 


6 


3 


— 


,, 


— 


61 


7 


3* 


18 


,, 


1259 


59 


7 


3i 


28 


,, 


A1485 


62 


7 


3jk 


30 


Southern Venturer 


1236 


74 


8 


4 


32 


Balaena 


— 


65 


9 


4* 


34 


,, 


1 745 A 


65 


9 


4* 


25 


Southern Venturer 


1239 


66 


9 


4* 


40 


Balaena 


A1261 


67 


10 


5 


40 


>> 


A1234 


67 


10 


5 


40 


,, 


— 


68 


16 


8 


35 


>t 


— 


70 


11 


5i 


— 


>> 


A1232 


70 


11 


5i 


40 


>> 


A1250 


70 


12 


6 


37 


>t 


A1550 


70 


12 


6 


— 


Southern Venturer 


1741 


68 


12 


6 


45 


tt a 


A1532 . 


74 


12 


6 


— 


Balaena 


A1255 


71 


12 


6 


— 


,, 


A1539 


72 


13 


<* 


40 


,, 


A1228 


72 


14 


7 


50 


,, 


A1233 


72 


15 


7* 


43 


„ 


— 


68 


16 


8 


95 


,, 


1251 


76 


18 


9 


50 


,, 


1253 


76 


19 


9i 


34 


,, 


A1486 


72 


20 


10 


58 


Southern Venturer 


— 


74 


21 


io£ 


55 


Balaena 


1252 


75 


21 


io£ 


80 


,, 


A1494 


72 


21 


io* 


58 


Southern Venturer 


A1240 


72 


22 


II 


78 


Balaena 


1247 


78 


22 


II 


90 


,, 


A1490 


75 


24 


12 


— 


Southern Venturer 


1230 


73 


28 


14 


— 


Balaena 


A1348 


77 


29 


I4i 


78 


,, 


Ai547 


75 


30 


15 


65 


,, 


1478 


73 


31 


I5i 


88 


Southern Venturer 


— 


75 


32 


16 


67 


Balaena 


A1235 


75 


32 


16 


85 


,, 


1747 


73 


32 


16 


55 


Southern Venturer 


A1242 


73 


33 


16J 


48 


Balaena 


— 


75 


34 


17 


100 


,, 


1742 


68 


38 


19 


85 


Southern Venturer 


— 


72 


42 


21 


105 


Balaena 


A1739 


75 


43 


21* 


64 


Southern Venturer 


A1536 


78 


44 


22 


60 


,, a 


A1538 


77 


46 


23 


60 


Balaena 


1243 


74 


47 


23i 


115 


,, 


A1748 


76 


48 


24 


64 


Southern Venturer 


A1740 


74 


50 


25 


87 


,1 a 


Ai547 


75 


30 


15 


65 


Balaena 


Ai555 


79 


53 


26£ 


75 


a 


Ai534 


75 


62 


31 


100 


a 


— 


78 


66 


33 


83 


a 



i6o EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 



Table D. — Male Antarctic Fin Whales Balaenoptera physalus Sample Area II 



Whale 


Total 


Number of 


Estimated 


Length of 




number 


length (ft.) 


laminae 


age (years) 


core (mm.) 


Factory ship 


A1487 


55 


5 


2j 


20 


Southern Venturer 


— 


63 


8 


4 


10 


Balaena 


1488 


63 


8 


4 


36 


Southern Venturer 


— 


63 


9 


4* 


35 


Balaena 


1497 


63 


9 


4* 


53 


Southern Venturer 


1416 


65 


9 


4* 


58 


„ 


A1749 


55 


9 


4* 


30 


tt > > 


A1492 


63 


10 


5 


53 


>t >t 


1475 


66 


11 


5* 


30 


tt it 


— 


65 


14 


7 


— 


Balaena 


— 


70 


16 


8 


50 


,, 


1744 


69 


16 


8 


24 


Southern Venturer 


— 


68 


19 


8* 


84 


Balaena 


1484 


7i 


19 


9* 


80 


Southern Venturer 


— 


71 


20 


10 


24 


Balaena 


— 


71 


20 


10 


62 


a 


— 


70 


22 


11 


50 


a 


— 


69 


23 


Hi 


35 


a 


1735 


69 


23 


Hi 


53 


Southern Venturer 


1737 


70 


3i 


i5i 


35 


>» It 


1495 


69 


30 


15 


63 


It It 


1489 


72 


32 


16 


50 


II II 


— 


68 


34 


17 


47 


Balaena 


— 


70 


34 


17 


— 


a 


— 


70 


36 


18 


30 


a 


— 


69 


40 


20 


75 


a 


— 


70 


40 


20 


43 


,, 


— 


70 


43 


21* 


40 


a 


— 


71 


44 


22 


24 


,, 


1736 


67 


45 


22^ 


95 


Southern Venturer 


A1488 


67 


50 


25 


36 


a a 


1481 


65 


59 


28i 


55 


' a >i 


A1738 


76 


63 


31* 


80 


a it 



EAR PLUG LAMINATIONS IN A POPULATION OF FIN WHALES 161 



Table E. — Unlogged Antarctic Fin Whales Balaenoptera physalus 



Males 



Females 



( 


> 


c 


> 




Estimated 




Estimated 


Whale 


age 


Whale 


age 


number 


(years) 


number 


(years) 


1600 


16 


1168 


4 


1321 


12 


773 


44 


i??5 


94 


474 


4 


890 


32^ 


757 


10* 


1715 


23i 


912 


44 


1969 


9 


1 164 


13 


— 


10 


1319 


14 


— 


17* 


480 


4 


— 


"4 


— 


9 


— 


8£ 


— 


i8£ 


— 


19 


— 


9 


— 


12 


— 


11 


— 


12* 


— 


94 


— 


12* 


— 


16 


— 


12* 


— 


6 


— 


44 


— 


64 





4 





8 
64 


— 


— 


— 


15 

74 




PLATE 5 

A series of ear plugs from female Fin Whales showing progressive lengthening of the core. 



Bull. B.M. (N.H.) ZooL, 5, 6. 



PLATE 5. 






PLATE 6 

A longitudinal section through the base of an abnormal ear plug showing coarse and fine 
laminations. Mag. x 20. 



Bull. B.M. (N.H.) Zool., 5, 6. 



PLATE 6. 




THE MONOTYPIC GENERA OF 
CICHLID FISHES IN LAKE VICTORIA 

PART II 

AND 

A REVISION OF THE LAKE 

VICTORIA HAPLOCHROMIS SPECIES 

(PISCES CICHLIDAE) 

PART III 



P. H. GREENWOOD 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 7 

LONDON: 1959 



THE MONOTYPIC GENERA OF CICHLID FISHES 

IN LAKE VICTORIA 

PART II 
AND 
A REVISION OF THE LAKE VICTORIA 
HAPLOCHROMIS SPECIES (PISCES CICHLIDAE) 

PART III 



BY 

P. H. GREENWOOD 

Department of Zoology, British Museum (Natural History) 



Pp. 163-218 ; 16 Text-figs. 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 7 

LONDON : 1959 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5, No. 7 of the Zoological 
series. 




Trustees of the British Museum, 1959 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued February, 1959 Price Sixteen Shillings 



THE MONOTYPIC GENERA OF CICHLID FISHES 
IN LAKE VICTORIA, PART II 1 

By P. H. GREENWOOD 2 

British Museum (Natural History), London 

CONTENTS 

Page 
Generic Diagnosis and Discussion . . . . . . .165 

Astatoreochromis alluaudi Pellegrin . . . . . . .167 

Description .......... 167 

Osteology .......... 169 

Affinities .......... 173 

Description and diagnosis of A . a. alluaudi Pellegrin and A . a. occi- 
dentalis subsp. nov. . . . . . . . .174 

Ecology 174 

Summary ............ 175 

Acknowledgments . . . . . . . . . -177 

References ........... 177 

A revision of the four endemic monotypic cichlid genera of Lake Victoria, Macro- 
pleurodus bicolor (Blgr.), Platytaeniodus degeni Blgr., Hoplotilapia retrodens Hilg., 
and Paralabidochromis victoriae Greenwood has already been published (Greenwood, 
1956). These species differ from Haplochromis in various dental characters. Unlike 
the other monotypic genera, Astatoreochromis alluaudi is not confined to the Lake 
Victoria basin ; its range includes Lakes Edward, George, Nakavali and Kachira 
(Trewavas, 1933). Furthermore, Astatoreochromis differs from Haplochromis only 
in having an increased number of spines in the anal fin ; the oral dentition is typically 
that of a non-piscivorous Haplochromis. 

Genus ASTATOREOCHROMIS Pellegrin, 1903 

Astatoreochromis Pellegrin, 1903, Mem. Soc. zool. France, 16, 385 ; Idem, 1905, ibid. 17, 185, 
pi. XVI, fig. 2 ; Idem, 1910, ibid. 22, 297 ; Regan, 1922, Proc. zool. Soc, London, 188 ; Fowler, 
1936, Proc. Acad. nat. Sci. Philad. 88, 333, fig. 138 (mis-spelt Astatore) ; Poll, 1939, Explor. 
Pare. Nat. Albert, mission H. Damas (1935-36), fasc. 6, 1-73. Haplochromis (part) Boulenger, 
1907, Fish, Nile, 505 pi. XC, fig. 4 ; Idem, 191 1, Ann. Mus. Genova (3), 5, 71 ; Idem, 1915, 
Cat. Afr. Fish. 3, 305, fig. 206. 

Type species. Astatoreochromis alluaudi Pellegrin, 1903. 

Diagnosis. Astatoreochromis differs from Haplochromis only in having four or 
more spines in the anal fin. From other genera in the Haplochromis group with more 
than four anal fin spines, Astatoreochromis is distinguished by the absence of a marked 
antero-posterior differentiation in the form of the premaxillary teeth. 

1 Part I was published in Bull. Br. Mus. nat. Hist., Zool. 3, No 7, 1956. 

2 Formerly East African Fisheries Research Organization, Jinja, Uganda. 

ZOOL. v, 7. 7 



1 66 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 

In comparison with the Haplochromis of Lakes Victoria, Edward, and Kachira, 
Astatoreochromis shows an increased ratio of spinous to branched rays in the dorsal 
and anal fins. From other Haplochromis-like genera in these lakes, Astatoreochromis 
differs both in having more anal fin spines and in the nature of its oral dentition. 

Discussion. As Boulenger (1907) pointed out, the principal diagnostic character 
for Astatoreochromis cannot be considered trenchant because some four-spined 
specimens of normally three-spined Haplochromis species have been recorded. He 
cites as an example an aberrant H. desfontainesi from Tunisia. Nevertheless, through- 
out the very numerous species of Haplochromis it is very exceptional to find an 
individual with more (or less) than three anal fin spines and as yet no specimens 
of Astatoreochromis with less than four anal spines have been found. It cannot be 
denied that Astatoreochromis and Haplochromis are closely related (as are Haplochromis 
and the other monotypic genera of Lakes Edward and Victoria) and it might seem 
that little is to be gained from recognizing Astatoreochromis as a distinct genus. 

However, Astatoreochromis differs from the Haplochromis of Lakes Victoria and 
Edward in four other characters which, if taken together, may indicate that it 
has a different lineage from these species. In an earlier paper (Greenwood, 1954) 
I drew attention to the form of the pharyngeal apophysis in Astatoreochromis and 
showed that it resembles the apophysis occurring in Haplochromis vanderhorsti 
Greenwood (Malagarasi River) and H. mahagiensis David & Poll (Lake Albert). 
The other Victoria species with enlarged pharyngeal bones (H. ishmaeli Blgr. and 
H. pharyngomylus Trewavas) have a different apophyseal form. A summary of 
these observations is given in Text-fig. 2. 

Contrary to my earlier views, I now consider that, taken by itself, apophyseal 
form is of doubtful value as an indicator of phyletic relationship. For example, 
both the H. mahagiensis-H . vanderhorsti and the H. ishmaeli-H. pharyngomylus types 
of apophysis are found in Lake Nyasa Haplochromis with enlarged pharyngeal 
bones ; Haplochromis placodon Regan (a species with hypertrophied pharyngeals) 
has the "ishmaeli" type whilst H. sphaerodon Regan, H. latristriga (Gunther) 
and H. selenurus (Regan) (species with less massive pharyngeals) have the " mahagi- 
ensis " type. There is no evidence to suggest that Nyasa fishes with " mahagiensis "- 
like apophyses are more closely related to one another than to H. placodon, or that 
they represent an exotic element within the Nyasa flock. Certainly there is no 
indication of their being related to the H. mahagiensis-H. vanderhorsti species group. 
Thus, one must conclude that similarity of apophyseal form is yet another example 
of convergent evolution, at least at an inter-group level. 

Considering Astatoreochromis in relation to the cichlid species flocks of Lakes 
Victoria and Edward it is clear that this genus does not conform to the general 
morphological pattern of the endemic species and genera. Three characters, the shape 
of the caudal fin, the coloration and the high number of anal ocelli, set Astatoreochromis 
apart. Excepting H. melanopterus (a species of doubtful validity, see Greenwood, 
p. 192) all the endemic Haplochromis of Lake Victoria have a truncate or subtruncate 
caudal fin ; the caudal of Astatoreochromis is distinctly rounded. 

A considerable variety of colour and colour patterns is exhibited by the endemic 



MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 167 

Haplochromis, but all can be broken down into various combinations of several 
basic types. The golden-green ground colour of Astatoreochromis does not occur 
in any endemic species. The third outstanding characteristic of Astatoreochromis 
is the high number of ocelli on the anal fin of male fishes. Not only are the ocelli 
more numerous than in Haplochromis, but they are arranged in three or four hori- 
zontal rows ; it is extremely rare to find more than two rows in any Haplochromis 
from Lake Victoria or Edward. 

In all these characters, Astatoreochromis resembles H. vanderhorsti. There is 
also one other point of close inter-specific resemblance ; both species show only 
slight dimorphism in the coloration of the two sexes. In contrast the coloration of 
Lake Victoria Haplochromis is markedly dimorphic. 

Thus, although the form of the pharyngeal apophysis alone is of doubtful value 
in showing phyletic relationships, I consider that the additional evidence supports 
my original conclusion that Astatoreochromis was derived from an H. vanderhorsti-like 
stem. The two other Victoria species with enlarged pharyngeal bones and dentition 
(H. ishmaeli and H. pharyngomylus) are apparently related to one another. Their 
origin was probably by way of two forms represented in the present lake by a genera- 
lized species formerly confused with H. michaeli [see Greenwood, 1954 and 1956a]., 
but now known to be an undescribed species and a species partly advanced towards 
extreme hypertrophy of the pharyngeal mill (H. obtusidens). 

The apparently distinct origin of Astatoreochromis alluaudi in relation to the rest 
of the Victoria-Edward Haplochromis species flock is a further and perhaps more 
fundamental reason for maintaining the species as a distinct genus. 



Astatoreochromis alluaudi Pellegrin, 1903 
(Text-fig. 1) 

For synonymy see under genus. 

Lectotype. A female 122 mm. S.L. from the Kavirondo Gulf, Lake Victoria ; 
Reg. No. 04, 137 of the Paris Museum. 

Description. From the available material it seems that only two characters 
(length of the caudal fin and the extent to which the lower pharyngeal bones are 
hypertrophied) show clear-cut differences between populations inhabiting the 
various lakes. These two characters will be treated separately but all others are 
given for the species as a whole. 

The general species description is based on the following material : Lakes Victoria 
and Kyoga (including the Victoria Nile), yy specimens, 20-163 mm. S.L. (of which 
40, including the four syntypes, were used in obtaining proportional measurements) : 
Lakes Edward and George n specimens 24-0-80-0 mm. S.L.; Lake Nakavali, 18 
specimens, 50-137 mm. S.L. (of which n were used for proportional measurements) ; 
Lake Kachira, three specimens 66-78 mm. S.L. 

Depth of body 33-8-43-3 per cent of standard length, length of head 32-1-40-0, 
mean (M) = 35 per cent. Dorsal head profile fairly steeply sloping, straight or some- 
what decurved, becoming concave in larger individuals, 



168 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 

Preorbital depth, showing slight positive allometry with standard length, 1 1-1-17-5 
(M = 15-0) per cent of head length, least interorbital width 25-2-31-7 (M = 28-3) 
per cent. Snout as broad as long, its length 25-0-33-3 (M = 29-2) per cent of head. 
Eye diameter shows negative allometry with standard length, being 31-5-23-2 
(M = 27-2) per cent of head in fishes 20-80 mm. S.L. and 24-3-18-8 (M = 22-1) 
per cent in larger individuals. Depth of cheek positively allometric with standard 
length ; 12-8-26-0 (M = 21-3) and 20-0-27-9 (M = 24-2) per cent of head in the two 
size groups mentioned above. 

Caudal peduncle 11-0-15-2 per cent of standard length, its length /depth ratio 
1-0-1-4 (modal range i-o-i-i) or, rarely, deeper than long. 




Fig. 1 . A statoreochromis alluaudi alluaudi (from Boulenger, Fishes of the Nile) . 



Mouth horizontal or slightly oblique. Jaws equal anteriorly or, occasionally, 
lower somewhat projecting ; posterior tip of the maxilla reaching or almost reaching 
the vertical to the anterior orbital margin. Lower jaw 35-0-45-3 (M = 40-0) per 
cent of head length and 1-3-2-0 (rarely) times as long as broad (modal range 1-5-1-6). 

Gill rakers short and stout ; 8 or 9 (occasionally 10, rarely 7) on the lower limb 
of the first gill-arch. 

Scales ctenoid ; lateral line with 30 (f.12), 31 (f.21), 32 (f.20) or 33 (f.2) scales ; 
cheek with 3 or 4 (occasionally 5) series ; 4 or 5 (occasionally 6) scales between the 
origin of the dorsal fin and the lateral line ; 4-6 (rarely 7) between the pectoral 
and pelvic fin bases. 

Fins. Dorsal with 23 (f.2), 24 (f.4), 25 (f.15), 26 (f.68), 27 (f.n) or 28 (f.i) rays, 
comprising 16 (f.5), 17 (f.16), 18 (f.59), 19 (f.20) or 20 (f.i) spinous and 7 or 8 (rarely 
9) branched rays. Anal fin with 11 (f.3), 12 (f.67), 13 (f.30) or 14 (f.2) rays comprising 



MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 169 

4 (f.28), 5 (f.63) or 6 (f.n) spinous and 7 or 8 (rarely 6 or 9) branched rays. Pectoral 
fin shorter than the head, 22-3-29-4 per cent of standard length. 

Caudal fin rounded, longer in fishes from Lakes Nakavali, Edward and George 
than in those from Lake Victoria ; namely : length of caudal fin in Victoria specimens 
(N = 41) 21-4-28-5 (Mean 24-3) per cent of standard length ; in Lake Nakavali 
fishes (N = 4) 24-0-31-6 (M = 27-4) per cent, and in Lake Edward fishes (N = 9), 
24-0-31-6 (M = 27-0). This fin was damaged in two of the three specimens from 
Lake Kachira. 

Pelvic fin with the first ray produced and extending to beyond the vent or as 
far as the spinous part of the anal fin. 

Teeth. Even in the smallest specimen examined, the most posterior teeth in the 
upper jaw were unicuspid. In fishes less than 100 mm. S.L., the anterior and lateral 
teeth of the upper jaw and the entire outer series of teeth in the lower jaw are unequally 
bicuspid and relatively stout. In larger specimens, the dentition is a mixture of 
weakly bicuspid and unicuspid teeth ; fishes over 140 mm. S.L. (and some smaller 
individuals) have only stout, unicuspid teeth in the outer series of both jaws. There 
are 28-56 (modal range 40-46) outer teeth in the upper jaw. 

The small, tricuspid or unicuspid inner teeth are arranged in one or two rows. 

Osteology. Vertebrae : 15 + 14 in the single specimen examined B.M. (N.H.) 
Reg. No. 1911.3.3.111, from Kakindu, Victoria Nile. 

N eurocranial apophysis for the upper pharyngeal bones. The form of this apophysis 
was mentioned in the discussion on generic characters. Since the apophysis is of 
importance in defining cichlid genera, its variation and the probable factors influenc- 
ing its variability in Astatoreochromis will be outlined briefly. 



pro 







boc. 



12 3 4 

Fig. 2. Semi-diagrammatic representation of the shape and proportions of elements 
contributing to the upper pharyngeal apophysis in : (1) young Astatoreochromis 
a. alluaudi ; (2) adult A . a. alluaudi ; (3) adult Haplochromis vanderhorsti ; (4) adult 
Haplochromis ishmaeli. Scale constant. 



Although the shape and proportions of elements contributing to the apophysis 
are affected by the relative size of the pharyngeal bones, the characteristic group 
facies (see p. 170) is developed even in the absence of markedly hypertrophied 
pharyngeals (Text-fig. 2, (i)). In A. alluaudi it appears that the extent to which 
the basioccipital facets are enlarged and expanded depends primarily on the relative 
hypertrophy of the pharyngeals, and secondarily on the size of the fish. Thus, in 



170 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 

two specimens from Lake Victoria, one, 73 mm. S.L. with weakly developed pharyn- 
geals, has proportionately smaller basioccipital facets than the other, 63 mm. S.L. 
and with enlarged pharyngeal bones and teeth (cf. Text-fig. 2 (i) and 2 (ii)). Likewise, 
fishes 70 mm., 76 mm., and 80 mm. S.L., from Lake Edward, and two specimens 
71 mm. and 82 mm. S.L. from Lake Nakavali all have weakly developed pharyngeals, 
and apophyses comparable with the 73 mm. fish mentioned above. In this size-range 
it would appear that the size of the pharyngeal bones is exerting full influence on 
apophyseal form. 

The effect of overall size is demonstrated in a fish 125 mm. S.L. from Lake Nakavali. 
In this specimen the pharyngeal bones are weak in comparison with those of a com- 
parable sized fish from Lake Victoria (cf. Text-fig. 3, lower row, left and right). 
Yet, the apophyseal form is similar in the two specimens except for a slightly 
smaller surface area in the Nakavali fish. 

Lower pharyngeal bone triangular. The form of this bone (which depends on the 
degree to which it is hypertrophied) and the nature of its teeth show a marked 
difference between fishes from Lake Victoria (including Kyoga) and those from the 
other lakes (see Text-fig. 3). When specimens of equal sizes from different lakes 
are compared it is immediately obvious that those from Lake Victoria have more 
massive bones with a greater proportion of molariform teeth. As far as can be 
determined from available material there is a little geographical variation of this 
character in fishes from Lakes Edward, George, Nakavali and Kachira. In all 
these populations the bone is clearly less massive than in Lake Victoria fishes and 
there are fewer molariform teeth. When present, such teeth are generally confined 
to the two median rows ; any enlarged teeth in the lateral series are usually cuspidate. 

The difference in pharyngeal bone size can be expressed quantitatively by using 
the ratio of head length to pharyngeal bone width (measured from tip to tip of the 
upper arms) ; it is, however, less impressive an indication of disparity in massiveness 
than an actual comparison of individual bones. The ratio for specimens from the 
various lakes is : Victoria (including Kyoga) ; 2-4-3-1 (Mean 2-7 ; 32 specimens 
examined) ; Nakavali : 2-6-3-6 (Mean 3-1 ; 16 specimens) ; Edward and George : 
2-8-3-6 (Mean 3-0 ; 10 specimens) ; Kachira : 2-7-3-1 (Mean 3-0 ; three specimens). 

As specimens of A . alluaudi from Lake Victoria cover a sufficiently wide size-range 
it is possible to determine ontogenetic changes in tooth form and in the proportions 
of the bone. In the smallest specimen (20 mm. S.L.) the two median tooth-rows are 
composed of enlarged but cuspidate teeth and the bone is relatively coarse (Text-fig. 
3 top row, left). With increasing size, the bone becomes proportionately stouter 
and the median teeth larger and blunter (Text-fig. 3 middle row, left), as do some 
of the teeth in the lateral rows. In the great majority of fishes over 60 mm. S.L., 
only the most lateral series of teeth, and those in the upper corners of the bone, 
remain slender and cuspidate. The number of such non-molariform teeth is even 
further reduced in fishes greater than 120 mm. S.L. Only seven of the 78 fishes 
examined had pharyngeal bones and dentition less hypertrophied than the modal 
condition for their respective size-groups. 

Ontogenetic changes are less marked in A. alluaudi from the western lakes. 
The impression gained from these specimens is that the pharyngeal bones, apart 



MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 171 






0-15 




0-25 







0-25 



0-25 







0-5 



05 



Fig. 3. Lower pharyngeal bones and teeth (lateral and occlusal views) of : Top row, 
left Astatoreochromis a. alluaudi 20 mm. S.L.; right, A. a. alluaudi 48 mm. S.L. Middle 
row, left, A. a. alluaudi 60 mm. S.L.; right, A. a. occidentalis (Lake Nakavali) 63 mm. 
S.L. Bottom row, left, A. a. alluaudi 120 mm. right, A. a, occidentalis (Lake Nakavali) 
123 mm. S.L. Scale in centimetres. 



172 MONOYTPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 

from their greater size, may be compared with those of 20-30 mm. A. alluaudi 
from Lake Victoria. 

Nothing is known about the epigenetics of A. alluaudi and little is known of 
the feeding habits of populations in lakes other than Victoria. It is therefore impos- 
sible to define the causal factors for the marked intra-specific, geographical difference 
in pharyngeal bones and teeth. 

In Lake Victoria, A. alluaudi feed almost exclusively on Mollusca (see below) 
and particularly on the thick-shelled Melanoides tuberculata. Considering the extreme 
plasticity of bone and its response to intermittent pressure (see Murray, 1932 ; 
Weinmann & Sichner, 1947) it seems probable that the effects of crushing such 
prey might produce an adaptational thickening and strengthening of the pharyngeals. 
In this way, any genetical tendency towards pharyngeal hypertrophy (as manifest 
in the relatively coarse lower pharyngeals of post-larval A. alluaudi) would be 
reinforced. If, on the other hand, in the western lakes the species is not predominant- 
ly a mollusc eater, the adaptational stimulus for increased bone size would be less, 
and the bones might be relatively weak. Finally, the possibility of inter-populational 
genetic differences cannot be discounted, especially since the various lakes are 
geographically isolated. 

Some data seem to add weight to the first, i.e. adaptational, hypothesis. The 
stomach and intestinal contents of 13 Lake Nakavali fishes have been examined ; 
of these, two were empty. Five of the remaining 11 fishes had fed on small cichlid 
fishes, and six on bottom debris (plant tissue) and insects (both adult and larval). 
Despite a careful search, no remains of Mollusca were identified. Admittedly, 
13 specimens do not constitute an adequate sample, but, if 13 Lake Victoria 
A. alluaudi in the same size-range were examined, every specimen with intestinal 
contents would have yielded remains of Mollusca. 

Likewise, in four A. alluaudi from Lake Edward and one from Lake George, the 
predominant food was insects, although three individuals had scanty remains of 
small Gastropoda in the intestines. The snails could not be identified, except in 
so far as they were not Melanoides sp. 

Coloration in life (known only from Lake Victoria). Sexual dimorphism is less 
marked in this species than in Haplochromis and the other monotypic genera. 
Females and immature males. Ground colour golden, overlain with olivaceous green, 
becoming yellow ventrally ; a dark band runs obliquely downwards through the 
eye and becomes continuous with the lachrymal stripe, which runs obliquely back- 
wards to the anterior tip of the preoperculum ; often another dark band along the 
vertical limb of the preoperculum. All median fins olivaceous-yellow, the dorsal 
and anal outlined in black ; caudal maculate. Pectoral fins hyaline ; pelvics faintly 
yellow or hyaline. 

Breeding males. Coloration essentially that of females except that the spinous 
dorsal is suffused with maroon, as is the entire anal fin, and the soft dorsal is densely 
spotted with maroon maculae. Anal fin with numerous yellow ocelli arranged in 
three or four vertical and the same number of horizontal rows. Pelvic fins black, 
the first ray pearly. Cephalic markings usually more intense than in females. 



MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 173 

Preserved material : Adult males. Ground colour greyish-brown to brown, lighter 
ventrally ; five or six dark transverse bars, often interrupted ventrally, on the 
flanks ; occasionally an interrupted mid-lateral stripe. Cephalic markings as described 
above. Soft dorsal fin and entire caudal maculate ; lappets of spinous dorsal, 
margin of soft dorsal and entire margin of anal fin black. Pelvics black laterally, 
the first ray pearly. Ocelli on anal fin dark grey. Females and immature males. 
Ground colour as in males but lighter. Soft dorsal and entire caudal weakly maculate 
or immaculate. Cephalic markings fainter than in males. Anal fin without ocelli, 
but in some individuals a few, small, light spots occur in the position of the ocelli. 
Pectoral and pelvic fins hyaline. 

Affinities. The relationship of Astatoreochromis alluaudi to the other monotypic 
genera of Lake Victoria and to certain species of Haplochromis was discussed above. 
It only remains to consider Regan's suggestion that A. alluaudi is " Near H. gestri, 
especially distinguished by the increased number of dorsal and anal spines and the 
large blunt pharyngeal teeth ". (Haplochromis gestri is a synonym of H. obesus 
(Blgr.) (seep. 182). 

With the information now available on the anatomy and ecology of both species, 
it is clear that A. alluaudi and H. obesus are not closely related. Haplochromis obesus 
belongs to a group of endemic Lake Victoria species which has developed the 
highly specialized habit of feeding on the embryos and larvae of other cichlid 
fishes (p. 187.) Astatoreochromis , on the other hand, possesses the potentialities 
for developing into a highly specialized mollusc-eater, although one subspecies is 
apparently a generalized bottom feeder. Besides the morphological differences 
noted by Regan, there are marked dissimilarities in the dentition and jaws of the 
two species. On the scale of divergence found in the Haplochromis and related 
species occurring in Lake Victoria, A. a. alluaudi and H. obesus must be placed in 
very distinct lineages. 

Differences in caudal fin length and the form of the pharyngeal bones are sufficiently 
well-marked to warrant the recognition of two subspecies of Astatoreochromis, one 
occurring in Lakes Victoria and Kyoga (including the Victoria Nile), and the 
other in Lakes Edward, George, Nakavali and Kachira, and in the Semliki River. 

Admittedly one of the characters distinguishing the two groups could be considered 
a response to environmental differences (see p. 172) . On the other hand, the importance 
of geographical isolation must be recognized. At present, and probably for a con- 
siderable period in the past, the western group of Lakes (Edward, Nakavali and 
Kachira) have been isolated from Lake Victoria by extensive papyrus-swamp 
divides on the interconnecting river systems (see Worthington, 1932). Likewise, 
Lakes Kachira and Nakavali are isolated from Lake Edward by intervening papyrus- 
swamps. Thus, although Astatoreochromis is relatively tolerant of papyrus-swamp 
conditions (see p. 174) the existence of such extensive swamp divides must consider- 
ably reduce any gene flow between the different lakes. Unfortunately, there is insuf- 
ficient material from Lakes Edward, Kachira and Nakavali to determine whether 
a distinct subspecies occurs in each lake. At present, therefore, only two subspecies 
can be recognized. 



174 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 

Astatoreochromis alluaudi alluaudi Pellegrin 

Diagnosis. Astatoreochromis a. alluaudi differs from the other subspecies in 
having a more massive lower pharyngeal bone with a greater number of molariform 
teeth, see Text-fig. 3 (ratio of head length to width of lower pharyngeal bone 
2*4-3-i, Mean 27), and in having a shorter caudal fin (21-4-28-5 [Mean 24-3] per 
cent of standard length). 

Other, ecological differences will be discussed below. 

Distribution. Lakes Victoria, Kyoga and the Victoria Nile. 

Astatoreochromis alluaudi occidentalis subsp. nov. 

Type specimen. A male, 125 +35-0 mm. long, B.M. (N.H.) Reg. No. 1933.2.23. 
146, collected by Worthington from Lake Nakavali. 

Diagnosis. Differs from the nominate subspecies in having a finer lower pharyngeal 
bone with fewer molariform teeth, see Text-fig. 3 (ratio of head length to width 
of lower pharyngeal bone 2-6-3-6, Mean 3-0) and in having a longer caudal fin 
(24-0-31-6, Mean 27-2 per cent of standard length). 

Distribution. Lakes Edward, George, Nakavali and Kachira ; the Semliki River 
above the rapids. 

Ecology. Habitat. A. a. alluaudi, unlike the majority of Haplochromis species in 
Lake Victoria, is not confined to any particular type of substrate. Indeed, in this 
lake the subspecies is ubiquitous in all areas where the water is less than 60 feet 
deep. There are also indications that in Lake Victoria A. a. alluaudi may extend 
into deeper water. Graham collected one specimen in surface nets set over 193 feet 
of water some distance off-shore (Station 71 ; o° 20 j' S., 33 i|' E.; in the collec- 
tions of E.A.F.R.O. there is one other specimen caught by nets set on the bottom at 
ca. 180 feet (o° 4' S., 33 14' E.). 

During rainy seasons, post-larval A. a. alluaudi have been found in pools and 
streams some distance inside papyrus-swamps. Larger young (40-50 mm. S.L.) 
enter small temporary streams when these are flowing into the lake. Neither the 
papyrus-swamp habitat nor that of temporary streams is occupied by endemic 
Haplochromis or related species. Young and adults of the widely-distributed, 
fluviatile-lacustrine species H. nubilus (Blgr.) and H. multicolor (Schoeller) do, 
however, live in such habitats. 

No habitat data are available for A. a. alluaudi in the Victoria Nile and Lake 
Kyoga, nor for A. a. occidentalis in any lake. Specimens of the latter have been 
collected from the Semliki River near its source in Lake Edward. 

Food. Astatoreochromis a. alluaudi (Lake Victoria). The stomach and intestinal 
contents of 40 fishes (48-163 mm. S.L.) from different localities clearly indicate that 
A. a. alluaudi feeds almost exclusively on Mollusca, especially Gastropoda. In 
most of the specimens examined, some insect larvae were also found ; but, both in 
volume and numbers, these represented only a small fraction of the ingested material. 
The very fragmentary nature of the shells found in the alimentary tract precluded 



MONOYTPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 175 

accurate identification of the mollusc species eaten. However, it seems most probable 
that the principal gastropod prey is Melanoides tuberculoid (Muller), and the chief 
lamellibranch, Corbicula sp. 

Astatoreochromis a. occidentalis. Lake Nakavali. Thirteen specimens 50-137 
mm. S.L. were examined ; two were empty. In the largest fish, the entire alimentary 
tract was filled with plant debris ; five specimens (79-123 mm. S.L.) each contained 
fragmentary remains of small cichlid fishes (probably Haplochromis), with, in two, 
a little plant debris and some insect remains. The five smaller fishes (50-72 mm.) 
contained fragmentary insect remains (especially larval and adult Diptera) and 
plant debris. 

Lake Edward. Only four specimens (62-76 mm. S.L.) were available for gut 
analysis ; three contained a few unidentifiable fragments of mollusc shells together 
with bottom debris and the fourth (71 mm. S.L.), mostly adult insects (Diptera) 
and the very fragmentary remains of a small fish. Although the mollusc fragments 
could not be identified positively they were not derived from Melanoides. 

Lake George. The alimentary tract of the single fish available (80 mm. S.L.) 
contained fragments of adult insects. 

Lake Kachira. The three specimens examined (66-78 mm. S.L.) were all from 
one station and contained only bottom debris and plant remains (including water-lily 
seeds) ; a few fragments of insects were found in the intestine of one individual. 

Breeding. Both subspecies of Astatoreochromis alluaudi are female mouth-brooders; 
exact spawning sites are not known. In Lake Victoria, males of A. a. alluaudi less 
than 100 mm. S.L. are immature but females are mature at about 95 mm. S.L. 
The three specimens of A. a. occidentalis from Lake Kachira (66-78 mm. S.L. 1 <J 
and 2 $) are all sexually active, thus suggesting that in this lake the subspecies 
reaches maturity at a smaller size than A. a. alluaudi in Lake Victoria. Little infor- 
mation was obtained on the size of sexually mature A. a. occidentalis in other lakes ; 
a brooding female 57 mm. S.L. from Lake Nakavali and a ripe female 62 mm. long 
from Lake Edward seem to indicate that in these lakes female A. a. occidentalis 
also mature at a smaller size than do the females of A . a. alluaudi in Lake Victoria. 
It is possible that differences in the feeding habits of the two subspecies may be 
primarily responsible for the smaller adult size of A. a. occidentalis. 

A marked disparity was noticed in the sex ratio of A. a. alluaudi from Lake 
Victoria and A. a. occidentalis from Lake Nakavali ; there is insufficient material 
to determine the sex ratio in other localities. Using only those specimens whose 
sex could be ascertained with certainty, the ratio is 16 $ : 46 (J in Lake Victoria, 
and 1 $ : 7 $ in Lake Nakavali. Reasons for this discrepancy are obscure but at 
least any bias introduced by collectors selecting brightly coloured males can be 
discounted ; both sexes are remarkably similar in colour. Furthermore, collections 
from Lake Victoria were made so as to eliminate this bias. 

SUMMARY 

1. The monotypic genus Astatoreochromis alluaudi is redescribed. 

2. The generic characters are discussed, particularly from the phylogenetic view- 
point. It is thought that A . a. alluaudi was not derived from the same stem as other 



176MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 

Victoria and Edward species with hypertrophied pharyngeal bones and teeth. 
By the same tokens, Astatoreochromis is not closely related to the other and endemic 
monotypic genera of the two lakes. The genus is apparently related to such fluviatile 
species as Haplochromis vanderhorsti (Malagarasi River system) and H. straeleni 
(Congo system). 

3. Two subspecific groups may be recognized, one from the Lake Victoria system 
and the other from lakes in western Uganda. These groups are given subspecific 
status, namely : Astatoreochromis a. alluaudi from Lakes Victoria and Kyoga, and 
the Victoria Nile ; and A. a. occidentalis from Lakes Edward, George, Nakavali 
and Kachira, and the Semliki River. 

4. The feeding habits of the two subspecies are described. 

Study Material and Distribution Records. 

Astatoreochromis a. alluaudi 

Museum and Reg. No. Locality Collector 

Kenya 
Paris Museum 

04,137 . Kavirondo Bay . Alluaud 

(Lectotype) 

04,138-9 . „ 

(Paratypes) 
B.M. (N.H.).— 1904.6.281 . „ 

(Paratype, presented by Paris Museum) 



B.M. (N.H.).- 


—1958.7.9.2 


Kisumu Harbour 
Uganda 


E.A.F.R.O 


B.M. (N.H.).- 


—1906.5.30.506-9 


Entebbe 


Degen 


,, 


1906.5.30.505 


Bunjako 


1, 


,, 


I907.5-7.73-76 


Buddu Coast 


Simon 


,, 


1911.3.3.112-3113 


Jinja, Ripon Falls 


Bayon 


,, 


I958.7.9.3-5 


Grant Bay 


E.A.F.R.O 


>> 


1958.7.9-6 


Karinya (near Jinja) 


,, 


,, 


1958. 7.9. 7- J 6 


Jinja 


»» 


,, 


1958. 7.9. 18-21 


Pilkington Bay 


>> 


>> 


1958.7.9.22 


Thruston Bay 


>> 


»> 


1958.7.9.23 


o° 4 'S., 33°i4'E. 


»> 


>> 


1958.7.9.24-37 


Entebbe Harbour 


>» 


>> 


1958.7.9.38 


Beach nr. Nasu Point 


>» 


> > 


I958.7.9.39-4 


Stream at Bugungu, 
Napoleon Gulf 


" 


>> 


1958.7.9.50 


Ekunu Bay 


,, 


>> 


1958.7-9.51-58 


Ramafuta Island 
Tanganyika 


" 


„ 


1958.7.9. 1 


Mwanza 


>> 


>» 


1958.7.9. 17 


Majita 


,, 


„ 


1958. 7. 9. 41-49 


Godziba Is. 


„ 



Lake Victoria, Locality Unknown 
1908.5. 19.51 . — 

1928.5.24.370-372 . — 



D. Radcliffe 
M. Graham 



MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 177 

Collector 



Museum and Reg. No. Locality 

Lake Kyoga and the Victoria Nile 
,, 1911.3.27.21 . Between Lake Kyoga and the 



1911.3.3.108 
1911.3.3.109-110 



Murchison Falls 

Bululo, Lake Kyoga 

Kakindu, Victoria Nile 



F. Melland 
Bayon 



Astatoreochromis a. occidentalis 
Lake Kachira 
B.M. (N.H.). — 1933. 2. 23. 160-162 



1933.2.23. 137-140 . 



1933-2.23. 141 



1933-2.23.142-159 



Lake Edward 



Lake George 



Lake Nakavali 



E. B. Worthington 



ACKNOWLEDGMENTS 

It is with great pleasure that I acknowledge my gratitude to Dr. Ethelwynn 
Trewavas for her helpful advice and criticism ; to the authorities of the Museum 
National d'Histoire naturelle, Paris for allowing me to examine Pellegrin's type 
specimens ; to Dr. M. Poll of the Musee Royal du Congo Beige, Tervueren, who 
placed at my disposal several specimens from the Semliki River, and to Dr. Denys 
W. Tucker for his helpful criticism of the manuscript. 



REFERENCES 
(Other than those given in full in the synonymy) 

Greenwood, P. H. 1954. On two cichlid fishes from the Malagarazi River (Tanganyika) etc. 

Ann. Mag. nat. Hist. (12) 7 : 401-414. 
1956. The monotypic genera of cichlid fishes in Lake Victoria. Bull. Br. Mus. nat. 

Hist., Zool. 3, No. 7. 
Murray, P. D. F. 1936. Bones. Cambridge. 

Regan, C. T. 1922. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. : 157-191. 
Trewavas, E. 1933. Scientific results of the Cambridge expedition to the East African 

lakes, 1930-1. II. The cichlid fishes. /. Linn. Soc. {Zool.) 38 : 308-341. 
Weinmann, J. P. & Sicher, H. 1947. Bone and Bones. Henry Kimpton, London. 
Worthington, E. B. 1932. A Report on the Fisheries of Uganda. Crown Agents, London. 



A REVISION OF THE LAKE VICTORIA 
HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE), 

PART III 



By P. H. GREENWOOD 

British Museum (Natural History) London 
CONTENTS 



Introduction ..... 
Haplochromis cronus sp. no v. . 
Haplochromis obesus (Boulenger) 
Haplochromis maxillaris Trewavas . 
Haplochromis melanopterus Trewavas 
Haplochromis parvidens (Boulenger) 
Haplochromis cryptodon sp. nov. 
Haplochromis microdon (Boulenger) . 

Discussion of the Seven Foregoing Species 
Haplochromis plagiodon Regan & Trewavas 
Haplochromis chilotes (Boulenger) 
Haplochromis chromogynos sp. nov. 
Haplochromis aelocephalus sp. nov. 

Summary ..... 

Acknowledgments 

References .... 



Page 

179 
180 
182 
189 
192 
194 
198 
200 
203 
205 
207 
212 
214 
218 
218 
218 



INTRODUCTION 

Five of the seven species described in the first part of this paper are known to feed 
almost exclusively on the embryos and larvae of other cichlid fishes, especially 
Haplochromis. Data on the food of the sixth species are very inadequate but are 
nevertheless indicative of similar habits. The seventh species is known from only a 
few specimens, but various morphological similarities between it and two other 
species of this group suggest embryo and larval fish-eating habits. 

If the species on which these fishes prey are mouth-brooders, it can be said that 
none of the young found in the stomachs of the predators was of a size at which it 
would normally have left the parental mouth. 

Despite identical feeding habits, the members of this species group are morphologi- 
cally heterogeneous and exhibit convergence only in a tendency for the teeth to be 
deeply embedded in the oral mucosa and in having capacious mouths. Furthermore, 
in most species there is a marked infra-specific variability in gross morphology, 
especially of the head. It seems that the group is of polyphyletic origin. 

zool. 5, 7. 8 



180 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

The four species dealt with in the second part of the paper are, morphologically 
speaking, somewhat isolated from the other Haplochromis of Lake Victoria. All 
are insectivores. 

Haplochromis cronus sp. nov. 
(Text-fig. i) 

Holotype. A female, 135 mm. standard length, from Buka Bay, Uganda. 

Description, based on eight specimens, including the holotype, 1 14-135 mm. 
standard length. 

Depth of body 39-5-43*5 per cent of standard length, length of head 30-3-34-6 
per cent. Dorsal head profile strongly curved, with a well-defined but localized 




Fig. 1. Haplochromis cronus ; holotype. Drawn by Miss D, Fitchew. 

swelling above the anterior part of the eye. Preorbital depth 167-18-2, mean (M) 
17-5 per cent of head length ; least interorbital width 31-9-35*5 (M = 33-1) per cent. 
Snout slightly broader than long, its length 31*6-35-7 (M = 33-7) per cent of head ; 
eye diameter 23-1-26-3 (M = 25-2), depth of cheek 29-3-34-2 (M = 30-1) per cent. 

Caudal peduncle 15-2-17-3 per cent of standard length, 1*1-1*3 (mode 1-3) times 
as long as deep. 

Jaws equal anteriorly ; lips thickened ; posterior tip of the maxilla not bullate 
and almost completely hidden beneath the preorbital, extending to the vertical 
through the anterior part of the eye. Lower jaw stout and deep, its length 29*3-34*2 
(M = 30-1) per cent of the head, 1-2-1-4 times as long as broad. 

Gill rakers stout ; 8-10 on the lower limb of the first arch. 

Scales ctenoid ; lateral line with 32 (f.5), 33 (f.i) or 34 (f.2) scales. Cheek with four 
or five series. Five to 8 scales between the dorsal fin origin and the lateral line ; 
8 or 9 between the pectoral and pelvic fin bases. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 181 

Fins. Dorsal with 24 (f.7) or 25 (f.i) rays, anal (damaged in one specimen) with 
12 (f.7), comprising XV or XVI, 9-10 and III, 9 spinous andj branched rays for the 
fins respectively. Pectoral fin shorter than the head ; pelvic fins with the first 
soft ray produced and extending to the vent in females and to the anal fin in males. 
Caudal truncate, the rays noticeably coarse ; densely scaled over about four-fifths 
of its length (a most unusual character in Lake Victoria Haplochromis species). 

Teeth. The outer row in both jaws is composed of unicuspid, fairly stout teeth, 
implanted vertically and not hidden by thickened oral mucosa ; there are 40-56 
teeth in this series of the upper jaw. The inner teeth are small and unicuspid, arranged 
in two rows (three in one specimen) in both jaws, and are separated from the outer 
series by a distinct space. 

Lower pharyngeal bone triangular, the dentigerous area about 1-3 times as broad as 
long ; the teeth are slender and cuspidate. 

Syncranium. The syncranium is noticeable for its short and deep neurocranium 
(comparable with H. obesus ; see p. 185) and for the stout but otherwise unspecialized 
dentary. These characters were determined from a radiograph B.M. (N.H.) Reg. 
No. 957 and the partial dissection of one specimen. 

Vertebrae : 13 -f- 16 in the single specimen radiographed. 

Coloration of preserved material : Adult females and sexually quiescent males. 
Ground colour dark golden above, lighter below, with traces of a golden-yellow 
flush on the operculum : a broad, mid-lateral stripe of variable depth and intensity 
crossed by four or five broad but faint transverse bars on the flanks ; a well-defined 
lachrymal stripe. Dorsal fin hyaline, with dark spots and bars on the soft part 
(probably deep red in life) ; caudal hyaline (densely maculate in males) ; proximal 
two-thirds of anal fin dark, remainder light ; pelvic fins dark (black laterally in 
males) . 

One of the three females available has a typical " bicolor" (piebald black and yellow) 
coloration, similar to that described in several other and apparently unrelated 
Haplochromis species and in two monotypic genera (Greenwood, 1957, and p. 213). 

Sexually active male. Dark brown above, sooty-grey below ; transverse and lateral 
stripes faint except at their junction mid-laterally. Dorsal fin dusky, the soft part 
maculate ; caudal dusky and densely maculate ; anal dark, except for its extreme 
tip and two colourless ocelli. Pelvic fins black on the lateral half and dusky mesially. 

Distribution. Known only from Lake Victoria. 

Ecology : Habitat. Five of the eight specimens are from an exposed beach habitat, 
one is from the sandy littoral of a sheltered gulf, one from the mud-bottom sublittoral 
of a sheltered bay and one from shallow water near a reed bed (no other data available) . 
In no locality was the water more than 20 feet deep. 

Food. Four specimens contained food in the stomach ; in each, only larval 
cichlid fishes were found (in three fishes these were identified as Haplochromis) ; 
the number of larvae in each fish was : 127 (ca. 11 mm. long) ; 50 (ca. 11 mm.) 
and 41 (ca. 11 mm.). The remains found in the fourth fish were too fragmentary 
to allow even an estimate of numbers. 

Breeding. Two females were found with, in one, larvae and in the other, newly 



182 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

hatched embryos in the buccal cavity. Since the ovarian condition of these fishes 
was clearly " spent " it can be assumed that the young were the fishes' own brood 
and not prey. 

Affinities. Haplochromis cronus belongs to the small group of deep-bodied, broad- 
headed Haplochromis whose adult size is ca. 100-140 mm. S.L. The specialized 
mollusc-eating species H. pharyngomylus Trewavas and H. ishmaeli Boulenger 
may be cited as examples of this morphotype. H. cronus differs from all other 
members of the group in having a densely and extensively scaled caudal fin. The 
species shows some affinity with H. obesus and H. maxillaris, forms which may have 
evolved independently from an H. cronus-like ancestor. 

Diagnosis. From other species with a similar gross morphology, H. cronus can 
be distinguished, primarily, by its almost completely scaled caudal fin (four-fifths 
scaled in H. cronus cf. two-thirds scaled in other species). The relatively large, 
completely exposed, caniniform and recurved teeth of H. cronus, together with an 
unmodified lower pharyngeal dentition and the presence of a supra-orbital swelling, 
also serve to distinguish H. cronus from other morphologically similar species. 

Study material and distribution records 



Museum and Reg. No. 


Locality 

Uganda 


Collector 


B.M. (N.H.).— 1958.1.16.85 


Buka Bay 


E.A.F.R.O 


(Holotype) 






B.M. (N.H.).— 1958. 1. 16. 86-89 


y, y> 


>> 


1958. 1. 16.90 


Napoleon Gulf near Jinja 


>> 


1958. 1. 16.91 


Pilkington Bay 
Kenya 


>> 


,, 1928.5.24.408 


Port Victoria 
(Graham's St. No. 84) 


M. Graham 



Haplochromis obesus (Boulenger) 1906 
(Text-figs. 2 and 3) 

Pelmatochromis obesus (part) Boulenger, 1906, Ann. Mag. nat. Hist. (17) 17, 447 (type specimen, 
by restriction [specimen figured in Fish. Nile], only) ; Idem, 1907, Fish. Nile, 491, pi. LXXXIX 
fig. 5 ; Idem, 1915, Cat. Afr. Fish. 3, 414, fig. 283. 

Lipochromis obesus (Boulenger), Regan, 1920, Ann. Mag. nat. Hist. (9) 5, 45 (foot-note). 

Haplochromis obesus (Boulenger), Regan, 1922, Proc. zool. Soc. London, 170. 

Paratilapia gestri Boulenger, ign,' Ann. Mus. Genova (3), 5, 67, pi. I, fig. 3. 

Paratilapia gestri (part, holotype only). Boulenger, 1915, Cat. Afr. Fish. 3, 318, fig. 211. 

Haplochromis gestri (part, holotype of P. gestri only), Regan, 1922, Proc. zool. Soc. London, 170. 



The union of H. obesus and H. gestri might be questioned if only the type specimens 
of the two species were available ; indeed, for a long time I thought that the species 
were distinct. However, after examining a large series of H. gestri-like specimens, 
I am forced to conclude that the type and unique specimen of H. obesus is merely 
an aberrant individual from a species whose modal morphotype is " gestri "-like. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 



:8 3 



In my opinion, the critical specific character-complex is the broad and stout lower 
jaw, combined with a relatively fine premaxilla and a stout, posteriorly bullate 
maxilla. These characters are easily verified in the type specimens of H. obesus 
and H. gestri, and have been further confirmed in a radiograph of the former. 

In its gross morphology and partly in its physiognomy, the type of H. obesus 
differs from all except one of the 46 specimens now referred to this species ; these 
differences may be partly attributable to post-mortem changes and poor preservation. 
In Boulenger's figure (reproduced here as Text-fig. 2) the mouth is shown as it appeared 
when closed artifically, with the result that the gape is very oblique. With the passage 
of time, the specimen has softened and it is now possible to close the mouth more 
easily. If this is done, it will be seen that the angle of the mouth is only slightly 




Fig. 2. Haplochromis obesus ; holotype (from Boulenger, Fishes of the Nile). 



oblique and that the dorsal head profile, although sloping steeply, is not so markedly 
concave as it appears in the figure. The lower jaw closes within the upper and only 
the anterior part of the maxilla is covered by the preorbital. Whether or not this is 
due to a natural deformity or to post-mortem distortion, I cannot say. 

Although, in appearance, the majority of specimens resemble the holotype of 
H. gestri, there are several others which depart from that mode but still retain the 
diagnostic dentary, upper jaw elements and dentition of the species. 

Description, based on 48 fishes (71-170 mm. S.L.) including the type, and the 
holotype of H. gestri. 

Depth of body 33*6-47-3 per cent of standard length ; length of head 30-3-35-9 
per cent. Physiognomy variable, the dorsal head profile straight or very slightly 
concave in the interorbital region, sloping steeply (Text-fig. 4) ; most fishes resemble 
the figured specimen. Preorbital depth 12-5-17-3 (M = 15-4) per cent of head length, 
least interorbital width 27-3-37-0 (M = 32-2) per cent. Snout 1-20-1-33 times as 
broad as long, its length 28-0-39-5 (M = 33-5) per cent of head ; eye diameter shows 
negative allometry with standard length, being 23-7-30-4 (M = 27-6) per cent in 



184 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

23 fishes less than 125 mm. S.L. and 20-5-27-9 (M = 23-6) per cent in 25 larger 
individuals ; depth of cheek 21-8-31-8 (M = 27-6) per cent. 

Caudal peduncle 12-8-17-9 per cent of standard length, its length 1-0-1-6 (modal 
range 1-0-1-2) times its depth. 




Fig. 3. Haplochromis obesus ; typical form (holotype of Paratilapia gestri) 
From Boulenger, Ann. Mus. Genova, 191 1. 






2 cm. 
Fig. 4. Haplochromis obesus ; individual variability of head profile. 



Mouth slightly oblique, maxilla bullate posteriorly and only partly covered by the 
preorbital, reaching the vertical through some part of anterior half of eye. Lips 
somewhat thickened ; jaws equal anteriorly, the length of the lower showing a 
positive but widely scattered allometry with standard length, 40-0-54*5 per cent 
of head length. Length /breadth ratio of the lower jaw 1-0-1-6 (mode 1-3). 

Gill rakers short and stout, 9 or 10 (rarely 8 or 11) on the lower part of the first 
arch. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 185 

Scales ctenoid ; lateral line with 29 (f.i), 31 (f.n), 32 (f.18), 33 (f.15) or 34 (f.2) 
scales ; cheek with 3 or 4 (rarely 2) series. Six or 7 (rarely 8) scales between the dorsal 
fin origin and the lateral line ; 6-8 between pectoral and pelvic fin bases. 

Fins. Dorsal with 24 (f.21), 25 (f.26) or 26 (f.i) rays, anal with 10 (f.2), n (f.23), 
12 (f.21) or 13 (f.i), comprising XV-XVI (rarely XVII), 8-10 and III, 8 or 9 (rarely 
10) spinous and branched rays for the fins respectively. Pectoral shorter than the 
head. Pelvic fins with the first soft ray produced, more particularly so in adult males. 
Caudal subtruncate or less commonly, obliquely truncate. 

Teeth. Both the inner and outer series of teeth are deeply embedded in the oral 
mucosa, so that only the tips protrude. 

Except for the smallest specimens, the outer teeth in both jaws are relatively 
stout and unicuspid with conical crowns. In small specimens most teeth are unequally 
bicuspid, or there may be an admixture of uni- and bicuspid forms. There are 34-52 
teeth in the outer series of the upper jaw. 

The shape of the teeth is variable ; in most specimens the anterior and some 
lateral outer teeth in the lower jaw have the crown bent so that its tip is directed 
anteriorly. In the upper jaw there is an admixture of such teeth with the more 
usual recurved and conical types. Teeth with anteriorly directed crowns are known 
only in H. obesus, H. maxillaris and H. melanopterus. 

The inner teeth are unicuspid and slenderly conical in fishes over 100 mm. S.L.; 
in smaller individuals there is a combination of unicuspid and weakly tricuspid 
teeth. Anteriorly in both jaws the inner teeth are arranged in one or, less commonly, 
two series ; the interspace between inner and outer teeth is greatly reduced or even 
absent. 

Lower pharyngeal bone short and broad, the dentigerous surface 1-2-1-6 times 
as broad as long ; pharyngeal teeth cuspidate and laterally compressed. 

Syncranium. The most outstanding skeletal characteristic of H. obesus is the broad 
and stout lower jaw (Text-fig. 5a). The " obesus "-type dentary is unique amongst 
the Haplochromis of Lake Victoria. When compared with one of the larger but 
generalized species the dentary of H. obesus is noticeably bullate in the region where 
each ramus divides into ascending and horizontal rami. 

Departure from a generalized Haplochromis condition is also seen in the maxilla, 
which is deeper and more bullate posteriorly. The neurocranium closely resembles 
that of Hoplotilapia retrodens Hilgendorf and Platytaeniodus degeni Boulenger, 
since the preorbital face is short and the supra-occipital crest deep (Greenwood, 
1956). This intergeneric convergence is probably associated with the relatively 
massive lower jaw of all three species. 

When compared with other Haplochromis (e.g. H. cronus and H. pharyngomylus) 
having approximately the same adult size and similar body-form, it is obvious 
that the mouth of H. obesus is more distensible and more protrusible. These factors 
may be associated with the specialized feeding habits of the species (see p. 204). 

Vertebrae : 13 + 16 and 13 + 15 (type H. obesus). 

Coloration in life : Adult males. Ground colour dark malachite green shading to 
silvery-blue ventrally ; a coppery sheen on the operculum, chest and belly ; a 



186 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

distinct dark mid-lateral stripe. Dorsal fin dark, with an overall pinkish flush ; 
caudal and anal fins dark, the latter with four or five orange-yellow ocelli arranged 
in one or, more frequently, two rows. Sexually quiescent males have a similar but 
less intense coloration. Adult females. Olivaceous-silver shading to silver ventrally ; 
a distinct, dark mid-lateral stripe. All fins hyaline ; in some individuals there are 
small yellow spots in the position of the anal ocelli in males. Some females show 
a typical " bicolor " black and silver (or yellow) piebald coloration. No estimate of 
the frequency of " bicolor " individuals can be made from the data available ; such 
females have, so far, only been found in the Napoleon Gulf, near Jinja, Uganda. 

A second atypical colour-form is also known. Fishes showing the extreme expres- 
sion of this coloration are uniformly black, but lighter (sooty) ventral coloration is 




Fig. 5 (a). Haplochromis obesus ; outline of dentary, ventral view. 

more usual. Unlike the " bicolor " pattern, the dark form is not sex-limited and is 
known to occur in several different areas of the lake. Furthermore, it shows some 
intergradation with the usual coloration, at least in males. 

Colour in preserved material : Both sexes. Ground colour golden to dark brown 
(adult males generally darkest) ; a well defined, dark, mid-lateral stripe and an 
ill-defined dorsal stripe following the contour of the upper lateral-line ; 5-9 vertical 
bars on the flanks and caudal peduncle ; often faint indications of a lachrymal 
stripe. Pelvic fins black in adult males, otherwise colourless, as are all other fins ; 
soft dorsal and the entire caudal weakly maculate. 

Distribution. Lake Victoria and Lake Kwania (Kyoga system). 

Ecology : Habitat. Haplochromis obesus is apparently restricted to water less than 
50 feet deep in the littoral and sublittoral zones of Lake Victoria. Most of the 
specimens were caught over a hard substrate (sand, shingle or rock) but two were 
caught over a soft mud bottom. In all probability, the distribution of H. obesus 
is closely linked with the spawning and brooding areas of the cichlid fishes on whose 
embryos and larvae it preys. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 187 

Food. Of the 73 specimens examined, 18 had food in the stomach. In every one 
of these fishes, only fish embryos or larvae were found ; with one exception (a small 
cyprinid fish) the prey could be identified as Cichlidae. A hundred embryos at the 
same stage of development were recorded from one stomach and in many others the 
embryos or larvae were all at the same developmental stage ; embryos at different 
ontogenetic stages were, however found in some individuals. 

The possibility that these stomach contents did not represent food but rather the 
fishes' own young accidently swallowed, can be overruled by the following 
considerations : a mixture of early and advanced ontogenetic stages was found in 
one stomach ; embryos and larvae were found in the stomachs of both male and female 
fishes and it is unknown amongst the Lake Victoria cichlids for both parents to 
share brooding duties ; early embryos were identified in the stomach contents of 




Fig. 5(b). Haplochromis parvidens ; outline of dentary, ventral view. 

an immature female ; and finally, personal observations show that it is unusual 
for a brooding female to swallow her brood when she is captured ; generally, the 
young are jettisoned. 

It is not known how H. obesus or the other larval fish-eating species obtain their 
prey. The question is complicated because the principal source of food for these 
species is the young of other cichlid fishes. Both species of Tilapia in Lake Victoria 
and all species of Haplochromis whose breeding habits are known are female mouth- 
brooders. Although late larval cichlids do leave the parental mouth, the earlier, 
non-free-swimming stages do not, except when the parent is so harrassed that it 
jettisons the brood. Unless a number of Haplochromis are not mouth-brooders, 
it seems that the larval and embryo fish-eating species employ some means of forcing 
the parent fish to abandon its brood. It may be added that there is no evidence to 
indicate that any Lake Victoria Haplochromis are not mouth-brooders. 

Breeding. A single brooding female was recorded : young removed from the buccal 
cavity were in the germ-ring stage of development. There is no sex-correlated size 
difference in adult fishes and sexual maturity is reached at a standard length of 
about 85 mm. 



188 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Affinities. Regan (1922) suggested that H. gestri (= H. obesus) was near Astatoreo- 
chromis alluaudi Pellegrin. With the additional information now available on both 
species, this opinion is no longer tenable. A. alluaudi is a specialized mollusc-eater 
with hypertrophied pharyngeal bones and the consequent modifications to the syn- 
cranial architecture (see Greenwood, 1954). Although H. obesus has a markedly 
modified lower jaw and somewhat atypical upper jaw features, it is more closely 
related to the generalized Haplochromis species. The relationships of A. alluaudi 
lie, apparently, with some of the semi-specialized fluviatile Haplochromis of the 
Malagarasi and Congo rivers (Greenwood op. cit. and p. 167). Any resemblance 
between A . alluaudi and H. obesus is entirely superficial and attributable to the stout 
bodies and broad heads of the two species. 

Perhaps the closest relatives of H. obesus are H. cronus and H. maxillaris, with 
which species it not only shows certain similarity in gross and detailed morphology, 
but it also shares the same food requirements. 

Diagnosis. The shape of the lower jaw (in which there is usually a predominance 
of unicuspid outer teeth with anteriorly directed crowns) is the most trenchantly 
diagnostic character. The deeply embedded teeth, together with certain morpho- 
metric characters of the head, serve to distinguish H. obesus from other Lake Victoria 
Haplochromis species. 



Study material and distribution records 



Museum 


and Reg. No. 


Locality 
Uganda 


Collector 


B.M. (N.H.).- 


—1906. 5. 30. 311 


Bunjako 


Degen. 


(Holotype Pelmatochromis obesus) 






Genoa Museu 


m 






(Holotype Paratilapia gestri) 


Jinja 


Bay on. 


B.M. (N.H.), 


— 1958. 1 . 16. 140 


Ekunu Bay 


E.A.F.R.O. 


>» 


1958. 1. 16. 141 


Buka Bay 


»> 


" 


1958. 1. 16.142 


Channel between Yempita and 
Busiri Isles, Buvuma Channel 


• >> 


» 


1958. 1. 16. 143-150 . 


Beach near Nasu Point, 
Buvuma Channel 


• 


» 


1958. 1. 16. 154-156 . 


SE. tip of Ramafuta Is., 
Buvuma Channel 


• >> 


,, 


1958. 1. 16. 157 


Karinya, Napoleon Gulf 


,, 


" 


1958.1.16.158-161 


Entebbe Harbour 
Kenya 


• >> 


>> 


1958.1.16.151-153 . 


Kisumu Harbour 


. 




Lake Victoria, Locality Unknown 




»> 


1958. 1. 16. 162-164 


— 


»» 


>> 


1928. 5. 24. 341-2 


Lake Kwania 


M. Graham. 


,, 


1929. 1.24.509 


— 


. E. B. Worthington 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 189 

Haplochromis maxillaris Trewavas, 1928 
(Text-fig. 6) 

Pelmatochromis microdon (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 441 ; Idem, 1915, 

Cat. Afr. Fish. 3, 412. 
Haplochromis microdon (Boulenger), (part), Regan, 1922, Proc. zool. Soc. London, 173. 
Haplochromis maxillaris Trewavas, 1928, Ann. Mag. nat. Hist. (10) 2, 94. 

Lectotype. A male 114 mm. standard length (B.M. (N.H.) Reg. No. 1928.5.24.486) 
from Emin Pasha Gulf, Tanganyika Territory (2 31J' S., 31 43^' E.), Michael 
Graham's station 227. 

Description, based on 58 specimens (including the types) 90-160 mm. S.L. 
Depth of body 32-0-42-8 per cent of standard length, length of head 30-0-34-8 
per cent. Physiognomy variable, its shape partly dependent on the angle of the mouth 




Fig. 6. Haplochromis maxillaris ; holotype. Drawn by Miss M. Fasken. 



and whether the lower jaw protrudes or not ; dorsal head profile concave (markedly 
so in a few specimens) and sloping at an angle of 40°-5o°. A few specimens bear a 
superficial resemblance to H. obesus, but despite this variability in gross morphology 
there is a distinct modal specific facies (see Text-fig. 6). 

Preorbital depth 11-4-16-3 (M = 13-9) per cent of head length, least interorbital 
width 22-6-31-3 (M = 26-5) per cent. Snout slightly broader than long, rarely as 
long as broad, its length 25-8-34-0 (M = 30-3) per cent of head. Eye diameter shows 
negative allometry with standard length : in 15 fishes 60-100 mm. S.L. it is 30-0-38-0 
(M = 33-2) per cent of head and in 43 larger individuals it is 25-0-31-4 (M = 27-3) 
per cent. Depth of cheek positively allometric with standard length : for the two 
size groups as above, 18-8-24-2 (M = 20-7) and 21-6-27-8 (M = 26-4) per cent 
head length. 

Caudal peduncle 12-6-18-5 per cent of standard length, 1-1-1-7 (modal range 
1-2-1-3) times as long as deep. 



igo A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Mouth distensible and usually somewhat oblique when closed, but horizontal 
in a few specimens. Maxilla partially hidden by the preorbital, its posterior tip 
bullate and reaching the vertical to the anterior part of the eye or even as far as the 
pupil. Lips thickened. Lower jaw usually projecting, but in a few fishes the jaws 
are equal anteriorly. Length of lower jaw shows positive allometry with standard 
length, in fishes 60-100 mm. S.L. it is 39*0-47-5 (M = 44-8) per cent of head length, 
and in larger fishes 46-5-56-0 (M = 50-0) per cent. Breadth of lower jaw contained 
1-3-2-2 (modal range 1-5-1-8) times in its length. 

Gill rakers short, 10 or 11 (rarely 9 or 12) on the lower part of the first gill- arch, 
the lower one or two rakers often greatly reduced. 

Scales ctenoid : lateral line with 29 (f.2), 30 (f.7), 31 (f.22), 32 (f.20), 33 (f.4) or 
34 (f.i) scales ; cheek with 2 or 3 series ; Si~7 scales between the dorsal fin origin 
and the lateral line, 5-7 (rarely 8) between the pectoral and pelvic fin bases. 

Fins. Dorsal with 24 (f.19), 25 (f.36) or 26 (f.3) rays, anal with n (f.18), 12 (f.37) 
or 13 (f.i), comprising XV-XVI (rarely XVII), 8-10 and III, 8 or 9 (rarely 10) 
spinous and branched rays for the fins respectively. Pectoral shorter than the 
head. First soft ray of the pelvic fin produced, extending to the vent in females 
and to the soft part of the anal fin in adult males. 

Teeth. The inner and outer rows of teeth in both jaws are deeply embedded in the 
thickened oral mucosa ; in many specimens the inner series are invisible without 
dissection. Furthermore, the outer teeth of the upper jaw. are covered by the thick- 
ened and inwardly curved margin of the lip. 

Fishes less than 80 mm. S.L. have small, weakly and unequally bicuspid outer teeth 
in both jaws. In larger fishes these teeth are also small, but stout and conical ; 
those in the upper jaw are recurved, whilst those in the lower jaw generally have the 
crown curved anteriorly or outwardly. Similar teeth are found in H. obesus, but are 
not the predominant form in that species. 

In the three skeletons examined there were 34, 36 and 40 outer teeth in the pre- 
maxilla. 

The inner teeth are weakly tricuspid in small fishes and unicuspid in larger indi- 
viduals ; arranged in one or, rarely, two series and separated from the outer row by 
a small interspace. Inner teeth in the upper jaw are slightly recurved and implanted 
so as to slope posteriorly ; those of the lower jaw are vertical or directed anteriorly. 

Lower pharyngeal bone triangular, the dentigerous area 1-0-1-4 times as broad as 
long ; teeth slender and cuspidate, those of the two median rows sometimes coarser. 

Syncranium. The dentary of H. maxillaris departs slightly from the generalized 
type. As in the dentary of H. obesus there is a pronounced lateral bullation of the 
area surrounding the bifurcation into ascending and horizontal rami. 

The premaxilla and maxilla are similar to those of H. obesus, except that the pre- 
maxillary teeth are restricted to the anterior and antero-lateral areas of the bone. 
The neurocranium is of a generalized Haplochromis type. 

Vertebrae : 13 -f- 16 and 12 + 16 in two skeletons. 

Coloration in life : Adult males. Ground colour dark blue-grey, lighter ventrally, 
with faint indications of darker transverse bars on the flanks. Dorsal fin dusky, with 



A REVISION OF THE LAKE VICTORIA H APLOCH RO M I S SPECIES 191 

maroon spots between the rays of the soft part ; lappets orange-red. Caudal and 
anal fins smoky-grey, the latter with three to five yellow ocelli arranged in either one 
or two rows. Pelvic fins dusky. Coloration in life of immature males unknown. 
Females. Silver-grey ground colour. Dorsal fin greyish ; anal and caudal fins 
similar but with a yellowish flush and, on the caudal, ill-defined, dark spots. Pelvic 
fins very faint yellow. Several dark, but faint, transverse bars may appear on the 
flanks immediately after death. 

Colour of preserved material : Males. Dark, some with an underlying silvery 
ground colour, others almost black. Seven or more transverse bands are sometimes 
visible on the flanks. Dorsal and caudal fins hyaline and maculate, or dusky ; anal 
hyaline or dusky. Pelvic fins black. Females. Ground colour yellowish-silver to 
brown ; some with seven or more transverse bars. All fins hyaline or somewhat 
dusky ; anal and caudal weakly maculate. 

Distribution. Known only from Lake Victoria. 

Ecology : Habitat. Haplochromis maxillaris is apparently restricted to water less 
than 30 feet deep, and particularly to the littoral and sublittoral zones of the lake. 
Most specimens were caught over a hard substrate (sand or shingle), but a few were 
recorded from mud substrates. Thus the habitat preferences of H. maxillaris are 
almost identical with those of H. obesus, the two species frequently being caught in 
the same gear. 

Food. Forty of the 118 individuals examined had identifiable food in the stomach. 
The smallest specimen (44 mm. S.L.) proved exceptional in that the stomach was 
filled with Copepoda and blue-green algae. All the remaining 39 fishes had eaten 
cichlid embryos or larvae. In some individuals, the entire stomach contents were of 
prey at uniform developmental stage, whilst in other fishes two or more stages 
(often as widely different as early cleavage embryos and late larval fishes) were 
present. Both sexes were represented amongst the fishes examined, which came 
from numerous localities. 

The remarkable similarity between the food of H. maxillaris and that of H. obesus 
is noteworthy. Again, it is difficult to imagine how the food is obtained if the species 
preys on mouth-brooding cichlids. 

Breeding. There is no information on any aspect of breeding behaviour in this 
species. All fishes below 100 mm. S.L. were immature ; it seems probable that sexual 
maturity is reached at a length of about 105 mm. Males and females attain the same 
adult size. 

Affinities. The distensible mouth, stout and posteriorly bullate maxilla, and the 
thickened lips of H. maxillaris all suggest affinity with H. obesus. Furthermore, 
conical outer teeth in which the crown is directed anteriorly or laterally are known 
only in these two species (and H. melanopterus , see below, p. 194). Certain specimens 
of both species show convergence in gross morphology, but the characteristic lower 
jaw of H. obesus usually allows for immediate identification. Apart from these few 
convergent individuals, the two species differ in certain morphometric details of the 
head and each has a distinctive modal facies. It is difficult to assess the phyletic 



192 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

significance of the resemblances and differences between H. obesus and H. maxillaris. 
The species could be equally well derived from a common stem or from unrelated 
ancestors within the Lake Victoria species flock. 



Study material and distribution records 
Museum and Reg. No. 



B.M. (N.H.).— 1928.5.24.486 

(Lectotype, H. maxillaris) 
1958. 1. 16. 182-184 
1958. 1. 16. 188 



Locality 
Tanganyika 

Emin Pasha Gulf 

Majita 
Mwanza, Capri Bay 



Collector 

M. Graham. 
E.A.F.R.O. 



1958.1.16.165-171 

1958. 1. 16. 172-179 

1958. 1. 16.180 

1958. 1. 16. 181 
1958. 1. 16. 185-187 
1958. 1. 16. 189 
1958. 1. 16. 190-193 
1958. 1. 16. 194-198 
1906. 5. 30. 310 
1958. 1. 16. 199 
1958. 1. 16.205-214 



Uganda 

Beach near Nasu Point, 

Buvuma Channel 

Ramafuta Is., Buvuma Channel 

Njoga, Williams Bay 

Buka Bay 

Beach near Hannington Bay 

Pilkington Bay 

Bukafu Bay 

Entebbe Harbour 

Entebbe 

Ekunu Bay 

Napoleon Gulf, near Jinja 



Degen. 
E.A.F.R.O. 



1958 .1.16. 200-204 



Kenya 

Kisumu Harbour 



Lake Victoria, Locality Unknown 
1928.5.24.480-485 . — 

(Syntypes, H. maxillaris) 



M. Graham. 



Haplochromis melanopterus Trewavas 1928 
(Text-fig. 7) 

H. melanopterus Trewavas, 1928, Ann. Mag. nat. Hist. (10) 2, 94. 

Discussion. This problematical species is known from a single specimen which had 
suffered some post-mortem distortion before preservation. Its status is, therefore, 
all the more difficult to decide. 

Superficially, H. melanopterus is most distinctive. The lower jaw (which closes 
entirely within the upper) is short, narrow and pointed anteriorly. The preorbital 
is very shallow so that the greater part of the maxilla is exposed when the mouth is 
shut. This latter character, together with the peculiar arrangement of the lower jaw 
in relation to the upper, may be an artefact of preservation and post-mortem distortion. 
The shallow preorbital and the short, pointed lower jaw cannot, however, be attri- 
buted to these causes. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 



93 



The dentition closely resembles that of H. maxillaris both in the form of the teeth 
and their restricted distribution on the premaxilla (see p. 190). The immediate 
question raised is, are not perhaps the shallow preorbital and the lower jaw size 
and shape the result of some ontogenetic disturbance in the development of an 
individual H. maxillaris ? The apparent distortion of the upper jaw might then be 
considered teratological rather than the result of post-mortem distortion. 

In general appearance, H. melanopterus is unlike H. maxillaris but as Trewavas 
noted in her original description of the species, it is nearer the H. maxillaris-obesus 
complex than any other species group. Thus, it is impossible to give an adequate 




Fig. 7. Haplochromis melanopterus ; holotype. Drawn by Miss M. Fasken. 

answer to the question posed above. I have decided to treat H. melanopterus as a 
distinct species mainly on the grounds that it is difficult to determine whether or 
not the peculiar jaws are a teratological feature, and because the nuchal and pectoral 
squamation of the type is manifestly smaller than in either H. obesus or H. maxillaris. 
Also, the rounded caudal fin is a most unusual feature in a Lake Victoria Haplochromis. 



Description, based on the holotype, an adult male 127 mm. S.L. 



Depth* 


Head* 


Po. % 


Io. % Snt. % Eye % 


Cheek % 


L-J- % 


C.P.* 


35*5 


33'5 


n-8 


27-0 30-6 28-2 

* Percentage standard length. 
% Percentage head-length. 


23'5 


37'6 


142 



Dorsal head profile very concave. Mouth probably oblique ; maxilla stout and 
bullate posteriorly, about three-quarters exposed even when the mouth is shut. 
In this specimen the mouth is open and the mandible lies horizontally, but the lateral 
limbs of the premaxilla and the maxillae are almost vertical. Lower jaw narrowing 
rapidly at a point almost half-way along its length ; greatest width contained 1-3 
times in the length ; the entire lower jaw closing within the upper ; lips thickened. 



194 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Dentition very similar to that of H. maxillaris. Outer teeth conical, those of the 
upper jaw recurved and restricted to the anterior and antero-lateral aspects of the 
premaxilla. Teeth in the lower jaw have the crown directed anteriorly or laterally ; 
the anterior teeth of this series are somewhat stouter than the equivalent teeth in 
H. maxillaris. Teeth of the inner series small and unicuspid, arranged in two irregular 
rows in each jaw. 

The oral mucosa appears to have shrunk ; consequently the outer teeth are more 
exposed than those of H. obesus or H. maxillaris, but the inner teeth are deeply 
embedded. 

Lower pharyngeal bone broken, but apparently similar to that of H. maxillaris ; 
pharyngeal teeth slender. 

Gill rakers moderately coarse ; ten on the lower part of the first gill-arch. 

Scales ctenoid ; 33 scales in the lateral line ; cheek with 3 or 4 series. Nuchal 
and pectoral scales small. Seven scales between the dorsal fin origin and the lateral 
line ; 9 between the pectoral and pelvic fin bases. 

Fins. Dorsal with XV, 8 rays, anal with III, 8. Pectoral very slightly shorter 
than the head ; pelvic fins with the first and second soft rays of about equal length, 
not quite reaching the anal fin. Caudal rounded. 

Vertebrae : 14 + 16 (from a radiograph, B.M. (N.H.) Reg. No. 955 a). 

Colour : Adult male. Brownish dorsally, brownish-silver on the flanks and belly. 
Dorsal, caudal and anal fins dusky, pelvics black. 

Ecology : Habitat. Smith Sound, Tanganyika Territory (2 33' S., 32 50' E.) in 
12 feet of water over a mud bottom (Graham, 1929). 
Food. The stomach is packed with early embryos of a cichlid fish. 

Affinities. Trewavas (1929) compared H. melanopterus with H. obesus (then 
known only from the holotype). Now that more material of H. obesus is available, 
the resemblance is found to be less marked. In some respects the morphology of 
the types is similar, but in the detailed structure of the head and dentition, 
H. melanopterus would seem to be more closely allied with H. maxillaris. It may yet 
prove to be merely a teratological specimen of that species. 

Haplochromis parvidens (Boulenger) 1911 
(Text-fig. 8) 

Paratilapia parvidens Boulenger, 191 1, Ann. Mus. Genova (3), 5, 65, pi. I, fig. 1 ; Idem, 1915, 

Cat. Afr. Fish. 3, 322 fig. 215. 
Haplochromis nigrescens (Pellegrin) (part, holotype of P. parvidens only), Regan 1922, Proc. 

zool. Soc. London, 172. 

Regan (1922) considered H. parvidens to be a synonym of H. nigrescens (Pellegrin) 
1909. I have re-examined the holotypes of both species and find that, although at 
first sight the species do resemble one another, the dentition and form of the lower 
jaw in H. parvidens is most distinctive. Additional material now available confirms 
and emphasizes these differences. The two species also differ in their feeding habits ; 
H. nigrescens is an insectivore and predator on small fishes, whilst H. parvidens 
is a specialized predator on embryo and larval fishes. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 195 

Haplochromis parvidens differs from the other larval and embryo fish-eating species 
in having a more slender and acutely pointed head, characters which typify the less 
specialized piscivorous predatory Haplochromis in Lake Victoria. The shape of the 
lower jaw is, however, unlike that of any predatory Haplochromis species (see 
Text-fig. 5B). 

Description, based on 32 fishes (including the holotype) 63-163 mm. S.L. 

Depth of body 33*3-38-2 per cent of standard length, length of head 33-3-37*5 
per cent. Physiognomy relatively constant, the dorsal head profile straight or 
gently concave, sloping at an angle of 30°-35°. Preorbital depth 15-9-20-5 (M = 18-6) 
per cent of head length ; least interorbital width 22-2-28-0 (M = 25-1) per cent. 
Snout length 1-2-1-33 times its breadth and 32-0-41-3 (M = 37-7) per cent of the head; 
eye diameter 20-3-27-2 (M = 23-0), depth of cheek 19-7-27-0 (M = 24*0) per cent. 




Fig. 8. Haplochromis parvidens ; holotype (from Boulenger, A nn. Mus. Genova, 191 1). 



Caudal peduncle 13-6-16-8 per cent of standard length, 1-1-1-5 times as long as 
deep (modal range 1-3-1-4 times). 

Mouth widely distensible and protractile, slightly oblique or horizontal when 
closed. Lips thickened. Lower jaw of a characteristic shape (Text-fig. 5B), somewhat 
rounded in cross-section and narrowing rapidly from a point about half-way along its 
length ; consequently, the anterior part closes within the upper jaw. Length of 
lower jaw 43*3-55*5 (M = 48-0) per cent of head length (showing a weak positive 
allometry with standard length) and 1-5-2-5 (modal range 1-9-2-1) times its width. 
Posterior tip of the maxilla bullate and almost completely hidden beneath the pre- 
orbital, usually not reaching the vertical to the anterior orbital margin, but extending 
to below the anterior part of the eye in a few specimens. 

Gill rakers. Nine to 11 (rarely 8 or 12, mode 10) on the lower part of the first gill- 
arch. 

Scales ctenoid ; lateral line with 30 (f.2), 31 (f.14), 32 (f.15) or 33 (f.i) scales. Cheek 
with 3 or 4 series ; 6 or 7 (rarely 5) scales between the origin of the dorsal fin and the 
lateral line ; 6 or 7 (less frequently 5) between the pectoral and pelvic fin bases. 
zool. 5, 7. 9 



196 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Fins. Dorsal with 23 (f.i), 24 (f.io), 25 (f.19) or 26 (f.2) rays, anal with n (f.9), 
12 (f.22) or 13 (f.i), comprising XV-XVI (rarely XIV), 9 or 10 (rarely 8) and III, 
8 or 9 (rarely 10) spinous and branched rays for the fins respectively. Pectoral 
fin shorter than the head. First soft ray of the pelvic fin produced and extending 
to the anal fin ; proportionately longer in adult males. 

Teeth. Both the inner and outer rows of teeth are deeply embedded in the oral 
epithelium, with the inner series often completely hidden. The outer teeth are mainly 
biscuspid in fishes 63-110 mm. S.L., with some unicuspids present in larger individuals. 
In fishes above this size, the outer teeth are predominantly unicuspid, relatively 
slender and recurved ; laterally placed teeth point inwards. In the three skeletons 
available, there are 50, 54 and 62 outer teeth in the upper jaw. 

Teeth in the inner series are small, slender and weakly tricuspid in fishes less than 
115 mm. S.L., but are unicuspid in larger individuals. In most fishes, the inner 
teeth are implanted almost horizontally, so that their crowns point backwards. 
One or two (rarely three) series of inner teeth occur in each jaw and are separated 
from the outer teeth by a small but distinct interspace. 

Lower pharyngeal bone with a triangular dentigerous surface, i- 1-1-2 times as 
broad as long or, rarely, somewhat wider. Lower pharyngeal teeth fine and cuspidate. 

Syncranium. The premaxilla and the dentary of H. parvidens are outstanding 
osteological characters. In combination they are diagnostic of the species. 

In H. parvidens, the premaxillary pedicels are as long as, or longer than the denti- 
gerous limb of the bone, whereas in the majority of Lake Victoria Haplochromis 
(including the large-mouthed species) the pedicels are shorter. 

The mandible has been described above ; its skeleton clearly shows the marked 
anterior narrowing and the peculiar lateral bullation of the area surrounding the 
bifurcation into ascending and horizontal rami (a similar swelling is also seen in 
H. obesus and H. maxillaris). 

The maxilla is strictly comparable with that of H. obesus and H. maxillaris, 
but the neurocranium differs in having an elongate preorbital face. The preorbital 
part of the skull is about one-third of the basilar length as compared with one-fifth 
to one-quarter in generalized Haplochromis. In this respect the neurocranium of 
H. parvidens resembles that of a small predatory Haplochromis such as H. nigrescens. 

Vertebrae : 13 + 16 in two skeletons examined. 

Coloration in life : Adult males. Ground colour dark blue-black dorsally, silvery- 
blue ventrally. Dorsal fin sooty, with orange-red lappets ; caudal sooty, the dorsal 
and ventral tips orange-red. Anal fin deep maroon, with two or three red ocelli. 
Pelvics black. Coloration of immature males unknown. Females. Ground colour an 
overall olivaceous-green, with faint indications of five to nine dark transverse bars. 
All fins olivaceous. 

Colour in preserved material : Adult and immature males. Dark brown, some 
adults almost black ; faint traces of five to nine transverse bars may be visible on 
the flanks and caudal peduncle. Dorsal fin dark, the soft part faintly maculate ; 
pelvics black ; anal and caudal fins dusky. Females. Golden-brown ground colora- 
tion, some faintly barred. All fins hyaline, the dorsal and caudal fins sometimes 
maculate. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 



197 



Distribution. Lake Victoria and Lake Salisbury (a single specimen in the collections 
of the Uganda Game and Fisheries Department, Entebbe). 

Ecology : Habitat. Like H. maxillaris and H. obesus, Haplochromis parvidens 
is apparently confined to littoral and sublittoral zones where the water is less than 
50 feet deep. Unlike the former species, however, H. parvidens is less closely restricted 
to a particular substrate. Although all three species have been caught in the same 
habitat, the available data suggest that H. parvidens may be the only member of this 
trophic group to occur commonly over a mud bottom. 

Food. Seventeen of the 60 specimens examined had food in the stomach ; of these, 
15 had eaten cichlid embryos or larvae. The stomachs of the two other fishes were 
filled with a fatty, yellow, yolk-like substance. 

Breeding. There is no information on the breeding habits of H. parvidens. Fishes 
less than 105 mm. S.L. are immature ; there is no apparent correlation between sex 
and maximum adult size. 

Affinities. Despite the deeply embedded teeth, long premaxillary pedicels and 
unusual lower jaw, there is an overall similarity between H. parvidens and some of 
the structurally less-specialized predatory Haplochromis. Haplochromis nigrescens, 
with which H. parvidens was previously synonymized, exemplifies this apparent 
relationship. There is also some similarity between H. parvidens and H. cryptodon, 
and more particularly with H. microdon. Haplochromis parvidens could have evolved 
from either an H. nigrescens-like stem or from a species resembling H. cryptodon. 



Study material and distribution records 




Museum and Reg. No. 


Locality 


Collector 






Uganda 




Genoa Museum. (Holotype) 


Ripon Falls, Jinja 


Bayon. 


B.M. (N.H.).- 


—I9II.3.3-33 


,, >> 


>> 


,, 


1958. 1. 16. 95 


Kaianje 


E.A.F.R.O 




, 


1958. 1. 16.96-98 


Entebbe Harbour 


,, 




, 


1958. 1. 99 


Busungwe Bay 


>> 




, 


1958. 1. 16.100 


Ekunu Bay 


i> 




, 


1958. 1. 16. IOI 


Napoleon Gulf, near Jinja 


it 




, 


1958. 1. 16. 108 


>» >» 


t> 




> 


1958. 1. 16. 130-139 


>> >> 


n 




, 


1958. 1 . 16. 107 


Macdonald Bay 


,, 




, 


1958. 1. 16. 109-113 


Pilkington Bay 


it 




, 


1958.1.16.114-115 


Njoga, Williams Bay 


it 




, 


1958. 1 . 16. 116-129 


Beach near Nasu Point, 


n 






Buvuma Channel 


it 






Kenya 




>> 


1958. 1. 16. 104-106 


Kisumu Harbour 
Tanganyika 


it 


, 


1 


1958. 1. 16.92-94 


Mwanza, Capri Bay 


it 



ig8 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Museum and Reg. No. Locality Collector 

Lake Victoria, Locality Unknown 

1958. 1. 16. 102-103 . — . 

1928.5.24. 112 . — . M.Graham. 

1928.5.24.399-400 . — . 

1 928. 5. 24. 401-402 . — . 

Haplochromis cryptodon sp. nov. 

Holotype. A male, 123 mm. standard length, from a beach near Nasu Point, 
Buvuma Channel, Uganda. 

Description, based on 31 specimens (including the holotype) 92-130 mm. standard 
length. 

Depth of body 27-5-35-6 per cent of standard length, length of head 30-3-34-9 
per cent. Physiognomy relatively uniform, the dorsal head profile straight and sloping 
at an angle of 35°-4o°. 

Preorbital depth 12-5-17-6 (M = 15-4) per cent of head length, least interorbital 
width 21-2-25-7 (M = 23-6) per cent. Snout as broad as long or very slightly broader, 
its length 27-5-34-2 (M = 31-3) per cent of head ; eye 23-1-29-4 (M = 25-8), depth 
of cheek 17-7-25-7 (M = 23-6) per cent. 

Caudal peduncle 14-7-17-7 per cent of standard length, 1-3-1-7 (modal range 
1-3-1-5) times as long as deep. 

(Four specimens [two from near Nasu Point, Buvuma Channel, Uganda, and two 
from Majita, Tanganyika Territory] differ in being noticeably more slender [depth 
27-5-31-0 per cent S.L.] and in having less steeply sloping heads. The two Uganda 
specimens also have a somewhat longer lower jaw [46-0 per cent head length] than 
is modal. In all other characters these specimens agree with the generality of indi- 
viduals. Since they are amongst the five smallest specimens available, it is possible 
that their divergent characters may be " juvenile "). 

Mouth slightly oblique and moderately distensible ; lips slightly thickened. Pos- 
terior tip of the maxilla somewhat bullate and reaching or almost reaching the vertical 
to the anterior orbital margin. Lower jaw with a tendency to narrow rather abruptly 
at about its mid-point, but not narrowing so markedly as in H. parvidens ; in 
some specimens (particularly individuals less than 100 mm. S.L.) this character is 
only visible after dissection. Length of lower jaw 39-2-46-5 ( M = 42-3) per cent of 
head length, 1-3-1-9 (modal range 1-5-1-6) times as long as broad. 

Gill rakers moderately slender, 10 or n (less frequently 9), on the lower limb of 
the anterior arch. 

Scales ctenoid ; lateral line with 30 (f.2), 31 (f.7), 32 (f.17), 33 (f.2) or 34 (f.2) 
scales. Cheek with 2 or 3 (rarely 4) series ; 5-7 scales between the lateral line and the 
origin of the dorsal fin ; 6-8 (rarely 9) between the pectoral and pelvic fin insertions. 

Fins. Dorsal with 23 (f.i), 24 (f.14) or 25 (f.16) rays, anal with 11 (f.7), 12 (f.23) or 
13 (f.i) comprising XV-XVI, 9 or 10 (rarely 8) and III, 8 or 9 (rarely 10) spinous 
and soft rays for the fins respectively. Pectoral fins shorter than the head. First 
pelvic ray produced and extending to the spinous part of the anal. 

Teeth. The inner and outer series of teeth are deeply embedded in the oral 
epithelium, so that only the tips of the outer teeth are visible. In specimens less 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 199 

than 100 mm. S.L. the outer teeth are weakly and unequally bicuspid. In larger 
fishes this row is composed of small, unicuspid and slightly recurved teeth. 

Teeth in the inner series are either unicuspid or weakly tricuspid, and are arranged 
in one or two rows. In fresh material it is usually impossible to see these teeth unless 
the oral mucosa is dissected away. 

Lower pharyngeal bone triangular, its dentigerous area 1-0-1-5 times as broad 
as long ; pharyngeal teeth slender and cuspidate. 

Syncranium. The syncranium of H. cryptodon resembles the generalized Haplo- 
chromis type, except that the maxilla is somewhat stouter and the dentary shows an 
incipient departure from the generalized condition towards that of H. parvidens 
(see p. 196 and text fig. 5B). 

Vertebrae : 14 + 16 in the single specimen examined (Radiograph, B.M. (N.H.) 
Reg. No. 958). 

Coloration in life : Adult and immature females. Ground colour dark green-brown 
shading to light gold ventrally. All fins hyaline. The live coloration of males is 
unknown. 

Colour of preserved material : Adult males. Ground colour dark gun-metal dorsally, 
shading to greyish-green on the flanks and ventral surfaces ; chest and branchiostegal 
membrane dusky ; faint traces of a coppery sheen on the operculum and flanks. 
Five to seven faint but dark transverse bars are visible on the flanks and caudal 
peduncle. Dorsal fin dusky ; caudal and anal fins hyaline, the latter with two to 
four dead- white ocelli ; pelvic fins black. Females. Ground colour light golden- 
yellow, slightly darker dorsally ; in some individuals there are faint traces of about 
five, broad, transverse bands on the flanks. One adult is of particular interest since 
it displays incipient male coloration ; the pelvic fins are dusky as are the chest 
and branchiostegal membrane. In addition there are traces of a coppery sheen on 
the operculum. 

Distribution. Habitat. H. cryptodon has been recorded from only three localities, 
namely, the Napoleon Gulf near Jinja, a beach near Nasu Point (Buvuma Channel) 
and a beach at Majita, Tanganyika Territory. The apparent absence of H. crytodon 
from other localities is difficult to explain since it was one of the more abundant 
species at the Nasu Point station and formed a regular element of the seine-net 
catches there. Perhaps it is significant that the majority of H. cryptodon caught 
at Nasu Point were brooding or " ripe " females, thus suggesting that the area is 
used as a breeding ground. If this is so, the species may normally occur in some other 
habitat which could not be fished by conventional gear. 

Food. Only one fish, a female from Nasu Point, had ingested material in the stomach 
and intestine. The stomach was packed with recently fertilized cichlid ova, whilst 
numerous small fish vertebrae were found in the posterior intestine. The stomach 
contents may have been the female's own brood, but the presence of larval fish 
vertebrae in the faeces cannot be explained on the same grounds. 

With such little positive evidence it is impossible to generalize on the feeding 
habits of H. cryptodon. But the single record of gut-contents, taken together with 
the jaw structure of this species, suggests a diet of embryo and larval fishes. 



200 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Breeding. H. cryptodon is a female mouth-brooder. The smallest individual 
caught, a female 92 mm. S.L., was sexually mature. Males and females reach the 
same adult size. 

Affinities. With the exception of H. cronus, H. cryptodon is the most generalized 
of the species referred to this trophic group. In structure and proportions it shows 
greater affinity with H. microdon and H. parvidens than with the H. obesus-H. 
maxillaris-H . cronus section of the group. H. cryptodon was probably evolved from 
the complex of piscivorous-insectivorous species which are not markedly differenti- 
ated (except for their larger size) from the generalized Haplochromis stock in Lake 
Victoria. 

Diagnosis. From the generality of Lake Victoria Haplochromis species, H. cryptodon 
may be distinguished by its distensible mouth and almost completely hidden dentition. 
From other species showing these characters, it is distinguished by the shape of the 
head (and particularly of the lower jaw) and an absence of teeth with anteriorly 
directed crowns. 

Study material and distribution records 

Museum and Reg. No. Locality Collector 

Uganda 

B.M. (N.H.).— 1958. 1. 16. 31 . Beach near Nasu Point, . E.A.F.R.O. 

(Holotype, H. cryptodon) Buvuma Channel 

1958. 1. 26. 32-33 . „ . 

1958. 1. 16.37-62 . ,, . 

„ 1958. 1. 16. 34 . Napoleon Gulf, near Jinj a 

Tanganyika 
1958. 1. 16.35-36 . Majita 



Haplochromis microdon (Boulenger) 1906 
(Text-fig. 9) 

Pelmatochromis microdon (part ; holotype only) Blgr., 1906, Ann. Mag. nat. Hist. (7) 17, 441 ; 

Idem, 1913, Cat. Afr. Fish. 3, 412, fig. 282. 

Haplochromis microdon (Blgr.), (part), Regan, 1922, Proc. Zool. Soc, London, 173. 

When redefining H. microdon Regan (1922) noted his belief that the type specimen 
had a malformed lower jaw (which did not bite against the upper) and that its small 
teeth were due to this malformation. In the light of additional specimens, I am 
unable to agree with Regan, and conclude that the shape of the lower jaw and its 
small teeth are, indeed, some of the diagnostic characters of the species. Conse- 
quently, I find that the other species which Regan referred to H. microdon can no 
longer be considered conspecific ; they will be dealt with in a subsequent paper. 

The peculiar lower jaw of H. microdon closely resembles that of H. parvidens 
but the two species differ in other osteological characters. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 201 

Description, based on eight specimens (including the holotype) 1 14-148 mm. S.L. 

Depth of body 33*1-37-6 per cent standard length, length of head 31-2-34-8 
per cent. Physiognomy variable and dependent on whether the dorsal head profile 
is moderately or strongly concave. Preorbital depth 14-6-18-8 (M = 17-1) per cent 
of head length, least interorbital width 24-6-29-6 (M = 26-6) per cent. Snout as 
long as broad or very slightly longer ; its length 30-8-34-5 (M = 32-5) per cent of 
head, eye diameter 24-0-28-2 (M = 25-5), depth of cheek 19-2-26-0 (M = 23-8) 
per cent. 

Caudal peduncle 15-3-17-0 per cent of standard length, 1-3-1-5 times as long as 
deep. 




Fig. 9. Haplochromis microdon ; holotype (from Boulenger, Fishes of the Nile). 



Mouth oblique, distensible and moderately protractile. Jaws equal anteriorly 
or the lower very slightly shorter ; lower jaw always closing within the upper, its 
length 43-5-48-0 (M = 46-2) per cent of head length, 1-6-2-3 (modal range 1-9-2-2) 
times as long as broad. Premaxillary pedicels shorter than the dentigerous limb. 
Posterior tip of the maxilla slightly bullate, partly hidden by the preorbital and extend- 
ing to the vertical to the anterior orbital margin, or almost so. 

[One specimen, an adult female 114-0 mm. S.L., from Pilkington Bay, is not 
included in the general description given above. Although it differs sufficiently from 
the other specimens to raise doubts as to its identity, I do not consider it to be a 
distinct species. The possibility of this fish being a hybrid between H. microdon 
and some other species (? H. parvidens) cannot, however, be excluded. 

The dentition and lower jaw are of the " microdon-parvidens " type, but the 
dentary is narrower anteriorly and it is shorter than even the extreme specimens 
of either species. These characters, coupled with the large eye and short snout 
give this fish an unusual appearance which departs from both the " microdon " 
and the "parvidens " types. The mouth is oblique and the premaxillary pedicels 
short, so that the sum of characters places the specimen nearest H. microdon. 



202 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

The principal morphometric characters are : 

Depth* Head* Po. % Io. % Snt. % Eye % Cheek % L.j. % C.P.* 

33*3 34-2 i8-o 25-2 29-5 30-8 20-5 41-0 15-8 

* Percentage S.L. 

% Percentage head-length.] 

Gill-rakers slender, 9-1 1 (mode n) on the lower part of the anterior arch. 

Scales ctenoid ; lateral line with 30 (f.2), 32 (f.4) or 33 (f.2) scales. 

Fins. Dorsal fin with 24 (f.4) or 25 (f.4) rays, anal with 10 (f.i), n (f.3) or 12 
(f.4), comprising XV-XVI, 9 or 10 and III, 7-9 spinous and branched rays for the 
fins respectively. Pectoral fin shorter than the head ; first soft pelvic ray produced, 
reaching to the vent in females and to the spinous anal fin in males. 

Teeth. All the teeth are so deeply embedded in the oral epithelium that they are 
invisible in fresh material. The teeth in the outer series of both jaws are small, 
unicuspid and slightly recurved. The single or rarely double row of unicuspid inner 
teeth is implanted at an acute angle. A distinct interspace separates the inner and 
outer series of teeth. 

Lower pharyngeal bone triangular, the dentigerous area slightly broader than long ; 
lower pharyngeal teeth slender and cuspidate. 

Syncranium (from a radiograph, B.M. (N.H.) Reg. No. 961, and a partial dissection) . 
The dentary is almost identical with that 0/ H. parvidens, but is more slender. 
The premaxilla is of a generalized type and does not have the elongated pedicels 
which characterize H. parvidens. The neurocranium is apparently intermediate 
between that of H. cryptodon and that of H. parvidens. 

Colour of preserved specimens : Adult males. Ground colour very dark brown, 
the ventral abdominal region lighter in one specimen ; five or six, broad and dark 
transverse bars visible on the flanks of some individuals. Branchiostegal membrane 
black. Dorsal, caudal, pelvic and anal fins dark, almost black, the anal with four, 
whitish ocelli. 

Adult females. Ground colour greyish-brown, darker dorsally ; very faint indica- 
tions of five or six broad transverse bars on the flanks. Dorsal, anal, and caudal 
fins hyaline, the last weakly maculate especially on the upper half ; pelvic fins black. 

Distribution. Lake Victoria. 

Ecology : Habitat. Seven of the eight fishes examined were from littoral zones of 
the lake, and were taken over both firm and soft substrates ; two were caught at 
a depth of 30-36 feet and the others in water 6-12 feet deep. The habitat of the 
eighth specimen is not known. 

Food unknown. Since the dentition and jaw structure of H. microdon so closely 
resemble those of H. parvidens, the feeding habits of the two species may well be 
similar. 

Breeding. No data are available ; the eight specimens are all adults. 

Affinities. The most obvious relative of H. microdon is H. parvidens ; the lower 
jaw in both species is of a type otherwise unrepresented in the Lake Victoria Haplo- 
chromis species flock. The dentary of H. microdon is, however, a more extreme 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 203 

modification of the generalized type than is the dentary of H. parvidens. On the 
other hand the elongated premaxillary pedicels of H. parvidens do not occur in 
H. microdon which retains a premaxilla of the generalized type. Consequently the 
mouth is less protrusible in this species. H. parvidens, in its gross morphology, and 
particularly in the shape of its neurocranium, shows strong affinity with some of the 
piscivorous predators (e.g. H. nigrescens). In corresponding characters, H. microdon 
is near H. cryptodon, from which it could be derived by further specialization of the 
lower jaw. Thus, the resemblance between H. microdon and H. parvidens may be 
the result of convergent evolutionary trends. 

Diagnosis. Haplochromis microdon may be distinguished from other species with 
deeply embedded teeth by the shape of the lower jaw, the concave dorsal head 
profile and the oblique mouth. It may be further distinguished from H. parvidens 
by its having premaxillary pedicels which are shorter than the horizontal (dentigerous) 
limb of this bone. 

Study material and distribution records 

Museum and Reg. No. Locality Collector 

Uganda 

B.M. (N.H.). — 1906.5.30.309 . Bunjako . Degen. 
(Holotype Pelmatochromis microdon) 

1958. 1. 16. 24 . Pilkington Bay . E.A.F.R.O. 

1958. 1. 16.25 • Entebbe Harbour 
,, 1958. 1. 16. 26 . Ekunu Bay 

,, 1958. 1 .16.29-30 . Beach near Nasu Point, 

Buvuma Channel 

Lake Victoria, Locality Unknown 
1958. 1. 16. 27-28 . — . 



DISCUSSION 

The embryo-larval fish-eating habits of this species-group were briefly mentioned 
in the introduction. As far as I can determine from published accounts, no other 
African cichlids have occupied this particular niche. 

That the group preys almost exclusively on the embryos and larvae of other 
cichlids is probably due to the fact that only the Cichlidae breed continuously ; 
the non-cichlid fishes spawn biannually, at the periods of maximum rainfall. On 
the other hand, the mouth-brooding habits of most Lake Victoria Haplochromis, 
and both Tilapia species, would seem to provide very little opportunity for these 
predators unless they have evolved a method of forcing the parent to jettison its 
brood. Since embryos at all stages of development have been found in the stomach 
contents, it is clear that the species do not obtain their food solely by preying on 
newly fertilized eggs before these are picked up by the brooding parent. The method 
of attack is unknown, but from the large number of embryos or larvae taken it must 
be highly efficient. 



2o 4 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Two anatomical features (a distensible and somewhat protrusible mouth, and a 
weak to moderately developed dentition deeply embedded in the thickened oral 
epithelium) characterize six of the seven species in this trophic group. It is not 
known if either of the characters has any functional significance in connection with 
the feeding habits. The seventh species, H. cronus, has strong and fully-exposed 
teeth, and the mouth is not noticeably distensible or protractile. Unfortunately, 
only a few specimens of H. cronus had food in the stomach. However, it may be signi- 
ficant that these fishes had fed on larger (ca. n mm.) late larvae, whilst members of 
the other species had fed on embryos and early larvae. That is to say, H. cronus 
had taken young of an age when they frequently leave the parent's mouth for short 
periods. Those species with distensible, protractile mouths and hidden teeth had 
taken mainly intra-oral young. Perhaps these larger mouthed species engulf the 
head or mouth of a brooding female and in this way force the parent to abandon its 
young? 

As in most other trophically defined species-groups there is evidence of both phyletic 
and convergent relationships between the species. But, unlike most others, this group 
shows greater divergence and more tenuous relationships within the apparently 
phyletic lines. 

Haplochromis cronus is the least specialized species, but it differs from the generality 
of Lake Victoria Haplochromis in having the caudal fin almost completely scaled 
(a characteristic of Lake Nyasa Haplochromis). 

Haplochromis obesus and H. maxillaris seem to be much specialized derivatives of 
a form resembling H. cronus, but neither of these species has a scaled caudal fin. 
The dentition is similar in H. obesus and H. maxillaris and quite unlike that of H. 
cronus ; in other characters (especially the shape of the lower jaw) the two species are 
markedly different. It is, in fact, impossible to decide whether the species are of the 
same lineage or the descendants of distinct but related ancestral stocks. 

A similar state of affairs exists when H. cryptodon and H. microdon are considered. 
A further complication is introduced by the resemblance between H. microdon 
and H. parvidens. In this case, however, it is possible that H. microdon was derived 
from an H. cryptodon-like ancestor, and H. parvidens from one of the less-specialized 
piscivorous predators. 

In all these species, anatomical differences between members of possible lineages 
are certainly greater than those encountered between species in the algal-grazing 
and mollusc-" shelling ' ' groups (Greenwood, 19566 and 1957). 

Two of the species described above, H. obesus and H. maxillaris, clearly demon- 
strate a phenomenon which is common amongst the Lake Victoria Haplochromis, 
namely, the intraspecific constancy of certain osteological and dental characters 
contrasting with the variability of other and often anatomically related characters. 
For example, the lower jaw and dentition of H. obesus is readily diagnostic, whilst 
the physiognomy is so variable that difficulty would be experienced in identifying 
some specimens were it not for the characteristic lower jaw. Likewise, there is marked 
variation in the gross morphology of H. maxillaris, yet the dentary and the dentition 
of both jaws are relatively constant. Haplochromis parvidens, however, shows only 
slight variation in its gross morphology. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 205 

In certain characters, Haplochromis taurinus Trewavas, of Lake Edward, resembles 
fishes of the H. maxillaris-H . obesus complex. The shape of the head approaches 
that of H. maxillaris, particularly with regard to the stout jaws and thickened lips ; 
also, the outer teeth in the upper jaw are hidden by a fold of lip-tissue. The likeness 
does not extend to the dentition, which is a critical character in this group. The teeth 
of H. taurinus are large, distinct and biscuspid (at least in the holotype, a fish 164 
mm. total length) and are of a form rarely encountered in any Lake Victoria Haplo- 
chromis. There can be little doubt therefore, that the resemblance between H. taurinus 
and H. maxillaris or H. obesus is superficial and of little phyletic significance. 



Haplochromis plagiodon Regan & Trewavas 1928 

(Text-fig. 10) 

Haplochromis crassilabris Blgr. (part), 1906, Ann. Mag. nat. Hist. (7) 17, 445. 

Paratilapia crassilabris (Blgr.), part, Blgr., 1915, Cat. Afr. Fish. 3, 345. 

Haplochromis crassilabris Blgr. (part), Regan, 1922, Proc. zool. Soc. London, 167. 

Hemitilapia bayoni Blgr. (part), Blgr., 1908, Ann. Mus. Genova (3), 4, 6 ; Idem, 1915, Cat. Afr. 

Fish. 3, 491. 
Clinodon bayoni (Blgr.) (part), Regan, 1920, Ann. Mag. nat. Hist. (9) 5, 45 (footnote). 
Haplochromis obliquidens Hilgendorf (part), Regan, 1922,- Proc. zool. Soc. London, 188. 
Haplochromis plagiodon Regan & Trewavas, 1928, Ann. Mag. nat. Hist. (10), 2, 224. 

Description, based on five specimens (including the holotype), 56-85 mm. standard 
length. 

H. plagiodon is a generalized species, except that it has teeth of an unusual form, 
resembling stout and somewhat modified versions of the teeth found in H. lividus 
Greenwood. Although H. plagiodon is represented by only five specimens, the form 
of the teeth is sufficiently distinctive and constant to warrant the assumption that 
the species is biologically valid. 

The principal morphometric characters for each of the five specimens are tabulated, 
below. All are males. 

S.L. Depth* Head* Po. % Io. % Snt. % Eye % Cheek % L.j. % C.P.* 



56-0 


35-7 


31-2 


I3-I 


25-7 


28-6 


30 -2 


21 -7 


34'2 


17-0 


72-0 


36-0 


3o-5 


15-0 


27-2 


27-2 


29-2 


22 -7 


36-3 


16-7 


76-0 


36-8 


3 i-6 


15-9 


25-0 


29-2 


3i-3 


23-0 


37'5 


15-2 


81-0 


32-7 


29-6 


16-7 


28-4 


29-2 


28-4 


20-8 


34-6 


18-0 


85-0 


36-5 


30-3 


15-6 


26-9 


26-9 


30-8 


23-0 


34-6 


16-5 



* Percentage standard length. 
% Percentage head length, 
f Holotype. 

Dorsal head profile straight and steeply sloping (ca. 50 ). Mouth horizontal ; posterior 
tip of the maxilla extending to the vertical to the anterior orbital margin, or slightly 
beyond. Jaws equal anteriorly, the lower 1-3-1-6 times as long as broad ; lips not 
thickened. 

Teeth. Outer teeth stout, erect and bicuspid, with the major cusp obliquely 
truncate and somewhat compressed, the minor cusp conical. In one specimen 



206 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

(72 mm. S.L.) the teeth are very worn, so that the minor cusp is indicated merely 
as a faint groove on the labial aspect of the tooth. There are 30-38 teeth in the outer 
series of the upper jaw. 

In most respects, the shape of these teeth closely resembles one of the variants 
occurring in H. lividus (see Text-fig. 2B in Greenwood, 19566), except that in 
H. plagiodon the teeth are not recurved, are stouter and do not have a distinct neck. 

The inner teeth are tricuspid and arranged in two or three rows in each jaw and 
are separated from the outer row by a distinct interspace. 

Lower pharyngeal bone triangular, fairly stout (as compared with, for example, 
fishes of the H. lividus-H. nuchisquamulatus group) ; dentigerous area 1 -2-1-5 




Fig. 10. Haplochromis plagiodon. Drawn by Miss D. Fitchew. 



times as broad as long. Lower pharyngeal teeth cuspidate, those of the two median 
series enlarged ; conical in three specimens but cuspidate in two others. 

Gill rakers short and stout ; 7 or 8 on the lower limb of the anterior arch. 

Scales ctenoid ; lateral line with 31 (f.i), 32 (f.3) or 33 (f.i) scales. Cheek with 
3 or 4 (rarely 2) series ; 6 or 7 scales between the dorsal fin origin and the lateral 
line, 8 or 9 between the pelvic and pectoral fin bases. 

Fins. Dorsal with 24 (f.i), 25 (f.2) or 26 (f.2) rays, anal with 11 (f.i), 12 (f.3) or 
13 (f.i) comprising XV-XVI, 9 or 10 and III, 8-10 spinous and branched rays for 
the fins respectively. First soft pelvic ray produced and extending to the spinous 
part of the anal fin. Pectoral fin slightly shorter than the head. 

Coloration. Unknown in life and known only for preserved males. Ground colour 
silver-grey to brownish-grey ; eight to ten transverse bars on the flanks and caudal 
peduncle, a fairly well-defined mid-lateral stripe, with indications of an interrupted 
band running slightly above the upper lateral line. Faint traces of two interocular 
bands and a lachrymal stripe. Dorsal, caudal and anal fins pale and immaculate ; 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 207 

six to eight small ocelli arranged in two rows on the anal fin. Pelvic fins dusky on 
the outer two-thirds in two specimens and entirely pale in the remainder. 

Ecology. No ecological data are available for the type and one other specimen; 
the three other fishes were caught in a seine-net operated in shallow water over an 
exposed, sandy beach at Entebbe, Uganda, In two of these specimens, remains of 
larval Diptera and Ephemeroptera (together with many fine sand grains) were found 
in the intestines. 

Diagnosis and affinities. Haplochromis plagiodon may be distinguished from other 
species in Lake Victoria by its peculiar teeth. The relatively coarse lower pharyngeal 
bone and the enlarged median pharyngeal teeth, together with the stout, firmly 
implanted and few (30-36) outer teeth, separate H. plagiodon from the two other 
species (H. lividus and H. nuchisquamulatus) with obliquely truncated, biscuspid 
outer teeth. 

Regan & Trewavas (1928) compared the teeth of H. plagiodon holotype with those 
of the type of Bayonia xenodonta Blgr. (now considered a synonym of Macropleurodus 
bicolor (Blgr.), Greenwood, 1956a). They emphasized the differences existing between 
the two species, even though there appeared to be some resemblance in the shape of 
the teeth. Now that additional specimens of comparable sizes are available for both 
species, it is clearer than ever that H. plagiodon is not closely related to Macro- 
pleurodus bicolor. Regan & Trewavas also suggested that H. plagiodon might be 
related to H. humilior (Blgr.). Although both these species have somewhat enlarged 
lower pharyngeal bones and median pharyngeal teeth, the resemblance in gross 
morphology is less marked and ceases when the oral dentition is compared. Likewise, 
although the oral teeth of H. plagiodon resemble those of H. lividus, the latter species 
has a fine lower pharyngeal bone with slender, cuspidate median teeth, and finer, 
more numerous oral teeth. 

Thus, it is only possible to suggest that H. plagiodon represents an independent 
offshoot from the generalized Haplochromis stem. 

Study material and distribution records 

Museum and Reg. No. Locality Collector 

Uganda 

B.M. (N.H.). — 1909.5.4.29 . Sesse Isles . Bayon. 
(Holotype H. plagiodon) 

1906.5.30.427 . Entebbe . Degen. 

1958. 1. 16.245-247 . Entebbe, Bugonga Beach . E.A.F.R.O. 

Haplochromis chilotes (Blgr.) 191 1 
(Text-fig. n) 

Paratilapia chilotes Blgr., 191 1, Ann. Mus. Genova (3), 5, 68, pi. II, fig. 2 ; Idem, Cat. Afr. 

Fish. 3, 338, fig. 228. 
Haplochromis chilotes (Blgr.), Regan, 1922, Proc. zool. Soc. London, 170. 

As defined below, H. chilotes exhibits a high degree of individual variability. The 
species may be divided into two morphotypes : first, those individuals with hyper- 



2o8 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

trophied and lobed lips and secondly, those in which the lips are thickened but not 
produced into well-defined lobes. Seventeen of the 25 specimens available fall into 
the first category and eight into the second. The division is not sharp, however, 
since the lips of some individuals in the second category do have a lobe-like, median 
swelling. 

Certain other characters are apparently correlated with the extent of lip develop- 
ment. For example, in most specimens with strongly lobed lips, the upper dental 
arcade is narrow and acutely rounded anteriorly. Also, in these fishes, the lower 
jaw is usually longer. The correlation is not complete and some lobe-lipped fishes 




Fig. 11. Haplochromis chilotes ; holotype (from Boulenger, Fishes of the Nile). 



have a mixture of these characters. It is because the sample as a whole shows a 
complete intergradation of lip, jaw and dental characters that I consider the material 
to represent a single species. Furthermore, I can discover no ecological differences 
between the two morphotypes, nor is there any obvious difference in the breeding 
coloration of the males. Nevertheless, the present arrangement should probably 
be considered tentative until more material and further field observations are available. 

Description, based on 25 specimens (including the holotype) 70-148 mm. standard 
length. 

Depth of body 32-5-40-8 per cent of standard length ; length of head (excluding 
the lips) apparently correlated with the degree of lip hypertrophy, 31-4-34-7 (M = 
32-7) per cent of standard length in fishes without clearly lobed lips and 32-0-38-2 
(M = 35-5) per cent in those with lobed lips. Dorsal head profile straight and gently 
sloping, or slightly decurved. Preorbital depth 15-0-18-2 (M = 16-8) per cent of 
head ; least interorbital width 19-3-27-2 (M = 23-8) ; snout longer than broad in 
lobe-lipped fishes and as broad as long in others, its length 30-8-38-4 (M = 34-0) per 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 209 

cent of head ; eye 28-6-21-8 (M = 25-4), depth of cheek 18-1-25-4 (M = 20-5) per 
cent. 

Caudal peduncle 12-5-17-2 (M = 15-3) per cent of standard length. 

Mouth horizontal, posterior tip of the maxilla reaching or almost reaching the 
vertical to the anterior orbital margin. Lips thickened, grossly so in some specimens 
in which each lip is produced medially into a tongue-shaped or globose lobe ; in 
others there may be an incipient lobe or even no indication of any such development. 
Jaws equal anteriorly, the lower proportionately longer (36-0-49-0, M = 39-6 per 
cent of head) in fishes with lobed lips than in the others (30-0-36-6, M = 33-2 per 
cent) . 

Gill-rakers short and stout, 7-9 on the lower limb of the anterior arch. 

Scales ctenoid ; lateral line with 31 (f.3), 32 (f.io), 33 (f.9), 34 (f.2) or 35 (f.i) 
scales. Cheek with 3 (rarely 2 or 4) series. Seven or 8 (rarely 6) scales between the 
origin of the dorsal fin and the lateral line ; 8 or 9 (less commonly 6, 7 or 10) between 
the pectoral and pelvic fin bases. 

Fins. Dorsal with 24 (f.3), 25 (f.18) or 26 (f.4) rays, anal with n (f.3) or 12 (f.22), 
comprising XV-XVI, 9 or 10 and III, 8 or 9 spinous and branched rays for the fins 
respectively. Pectoral fin shorter than the head ; first soft pelvic ray produced, 
variable in its posterior extension but proportionately longer in adult males than 
females. Caudal subtruncate. 

Teeth. In most specimens with hypertrophied and lobed lips the outer teeth in 
the upper jaw are arranged in an acutely rounded arcade, a pattern not found in any 
other Haplochromis from Lake Victoria. Specimens with thickened but non-lobed lips 
have more broadly rounded dental arcades, which are, nevertheless, more acutely 
rounded anteriorly than those of most other species. Complete intergradation 
exists between the various arcade shapes ; the correlation between lip development 
and dental pattern is not complete since some fishes with lobed lips have a broad 
arcade. 

In fishes 70-108 mm. standard length the outer tooth row is composed of unicuspid 
and weakly bicuspid, slender and slightly recurved teeth. In larger individuals 
these teeth are always unicuspid, are stouter and very slightly, if at all, curved. 
From 16 to 46 teeth may occur in the outer series of the upper jaw ; there is apparently 
some correlation between lip development and the number of teeth, with a tendency 
for lobe-lipped fishes to have more teeth. 

The inner series are composed of tricuspid and weakly tricuspid teeth in fishes less 
than no mm. standard length, and of predominantly unicuspid teeth in larger 
individuals. These teeth are arranged in two or three rows (less commonly four or 
one) anteriorly in each jaw. 

Lower pharyngeal bone triangular, the dentigerous area 1-0-1-5 (modal range 
1-2-1-3) times as broad as long. With one exception, all specimens have the median 
teeth (particularly the posterior few pairs) enlarged but still clearly cuspidate. In 
the exceptional specimen, the median teeth are slender. 

Coloration in life : Breeding males. Ground colour greyish-black or black ; lips 
and branchiostegal membrane black. Dorsal fin black, lappets and margin of soft part 



210 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

red, as are the maculae between the branched rays. Caudal black basally, but light 
yellow or hyaline distally ; anal yellow or hyaline, with three or four reddish-yellow 
ocelli ; pelvics black. Non-breeding males : Ground colour variable, usually greyish- 
black ; a dark mid-lateral band and a less intense, interrupted, wavy dorsal band, 
are generally visible. Dorsal, caudal and anal fins greyish ; pelvics black. 

Females and immature males. Ground colour greyish-silver, creamy- white on 
the chest and ventral surface of the head. Lips grey or cream ; six to nine dark 
transverse bars on the flanks and caudal peduncle, which are broadest at the points 
of intersection with the broad, mid-lateral stripe and the narrower, interrupted 
dorsal band. Dorsal caudal and anal fins greyish ; pelvics hyaline. 

One of the eight females available was a piebald, black and yellow " bicolor " 
variant (p. 212) ; it was of the group with hypertrophied and lobed lips. 

Preserved material : Sexually active males. Ground colour blackish, lower jaw, 
branchiostegal membrane and chest dusky ; lips usually pale but dusky in a few 
specimens. A dark, mid-lateral stripe and a less intense dorsal band, together with 
six or seven broad transverse bars, are usually visible on the flanks. Dorsal fin 
black, with light lappets and margin to the soft part. Anal fin pale orange-yellow, 
with three or four white ocelli arranged in a single row. Caudal fin black on the basal 
third to half, orange-yellow distally. Pelvic fins black, somewhat lighter medially. 
Sexually inactive males have a similar coloration except that the ground colour is 
lighter, as is the lower jaw and the branchiostegal membrane, whilst the chest is 
silvery. Females and juvenile males are brownish-fawn, lightest ventrally ; the 
banding and barring is more obvious than in adult males. All fins hyaline, with the 
soft part of the dorsal fin and the caudal fin maculate. 

Distribution. Known from Lake Victoria and probably also from the Victoria 
Nile since the type locality is given as " Victoria Nile at Ripon Falls " (Boulenger, 
1911). 

Ecology : Habitat. H. chilotes is apparently confined to the littoral and sublittoral 
zones of the lake, where the depth of water is less than 50 feet and to localities with 
a hard substrate (sand, shingle and rocks) . Only two specimens were caught over a 
mud bottom. Furthermore, it seems that the species may be confined to sheltered 
bays and gulfs ; with two exceptions, no H. chilotes have been recorded from exposed 
habitats. The exceptional fishes were caught near islands lying some distance from 
the mainland. 

Food. Eight of the 23 specimens contained food in the stomach and intestines ; 
these fishes represent five different localities, four of which are geographically distant 
from one another. 

One fish had fed almost exclusively on prawns (Caridina nilotica Roux) and the 
others on insect larvae. Larvae of the boring may-fly (Povilla adusta Navas) formed 
the main insect prey, and it was noticed that the silk case of the larva had also 
been ingested. Other insects eaten included Trichoptera and Dipt era larvae. From 
the amount of plant debris and sand occurring in the stomach it would seem that 
H. chilotes feeds from the bottom. 

Breeding habits. Females 70 mm. S.L. are sexually active, but the smallest adult 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 211 

male in this sample was 97 mm. S.L.; from the available data it seems probable that 
males reach a greater adult size than females. One female was found with three 
late larvae in the buccal cavity ; since the condition of this fish was clearly " spent " 
it is assumed that the larvae were part of a larger brood which was lost when the 
female was captured. 

Morphology of late larval H. chilotes. The three young fishes (all ca. 10 mm. S.L.) 
referred to above are indistinguishable from the larvae of other and unrelated 
Haplochromis species. Although the female parent had hypertrophied and lobed 
lips, no trace of these characters was visible in the larvae. 

Affinities. Haplochromis chilotes was probably derived from an H. chromogynos- 
like ancestor by further development of the lips, narrowing of the head and the conse- 
quent effects on the dental pattern. H. chilotes without lobed lips resemble H. chromo- 
gynos more closely than do specimens with lobed lips. But the similarity between 
the two species, even at its closest, is less marked than for example, that between 
H. sauvagei and H. prodromus (Greenwood, 1957). 

There is a striking superficial resemblance between H. chilotes and Lobochilotes 
labiatus (Blgr.) of Lake Tanganyika. The range of lip development in L. labiatus 
is about equal to that of H. chilotes but with this difference ; it is positively correlated 
with size in L. labiatus. Small fishes have weakly lobed lips whilst, in larger individuals 
the lobes are well developed. Tooth form in Lobochilotes is quite unlike that of 
H. chilotes, but there is a tendency for the dental arcade to be acutely rounded 
anteriorly. 

Lobed lips are also developed in Haplochromis lobochilotes of Lake Nyasa and thus 
there is some resemblance between this species and H. chilotes. In this instance, 
however, the similarity is less marked because the general proportions of the two 
species are somewhat different ; again, the form of the teeth is dissimilar. 

Of the lobe-lipped species occurring outside Lake Victoria, Melanochromis labrosus 
Trewavas, of Lake Nyasa shows the greatest overall and detailed likeness with 
H. chilotes. The dentition of M. labrosus and H. chilotes of a comparable size is similar, 
as is the gross and finer morphology of the head. The nuchal and pectoral squamation 
of M. labrosus is, however, markedly smaller than that of H. chilotes. Unfortunately, 
M. labrosus is known only from one specimen so a detailed comparison of the two 
species cannot be made. 

Diagnosis. H. chilotes with hypertrophied and lobed lips are immediately distin- 
guishable from other Lake Victoria species on this character alone ; from H. lobo- 
chilotes of Lake Nyasa and Lobochilotes labiatus of Lake Tanganyika, it is distinguished 
by differences in the shape of the teeth and by various morphometric characters 
Haplochromis chilotes with moderately developed and weakly or non-lobed lips may 
be confused with H. chromogynos or with species of the H. sauvagei complex. They 
are immediately distinguishable from the latter group in having fewer rows of inner 
teeth and by the shape of the outer teeth, which do not have strongly recurved tips. 
From H. chromogynos, H. chilotes is distinguished by its narrower interorbital region 
and longer snout. 

zool. 5, 7. 10 



212 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 
Study material and distribution records 



Museum and Reg. No. 


Locality 


Collector 








Uganda 




Genoa Museum 


Jinja, Ripon Falls 


Bayon. 




(Type Paratilapia chilotes) 






B.M. (N 


.H.), 


— 1911.3.3.33 


»» >, 


it 




(Paratype P. chilotes) 






,, 




1958. 1. 16.3 


Ramafuta Is. 


E.A.F.R.O. 


,, 




1958. 1. 16.4 


Pilkington Bay 


,, 


,, 




1958. 1. 16. 5 


Lukula Is. 


tt 


>> 




1958. 1. 16.6 


Channel between Dagusi Is. 
and mainland 


>t 


> > 




1958. 1. 16.9 
1958. 1. 16. 10-16 


Ekunu Bay 
Off south tip of Buvuma Is. 


tt 

1 1 


" 




1958. 1. 16. 17-23 


Napoleon Gulf, near Jinja 
Tanganyika 


II 


,, 




1958. 1. 16. 1 


Kaseiraji Is. 


II 


" 




1958. 1. 16. 8 


Godziba Is. 
Kenya 


tt 


» 




1958. 1. 16.2 


Kisumu Harbour 


» 






Lake Victoria, Locality Unknown 




>» 




1958. 1. 16.7 


— 


It 



Haplochromis chromogynos sp. nov. 

Haplochromis bicolor Blgr. (part), 1906, Ann. Mag. nat. Hist. (7) 17, 444. 
Paratilapia bicolor (Blgr.) (part), Blgr., 1915, Cat. Afr. Fish. 3, 346. 
Haplochromis sauvagei (Pfeff.), (part), Regan, 1922, Proc. zool. Soc. London, 167. 

Haplochromis chromogynos is unique amongst the Haplochromis of Lake Victoria 
(and probably the other lakes as well) since a high percentage, if not all, of the females 
have a piebald, black and yellow " bicolor" coloration. "Bicolor" female variants 
are known to occur in several Haplochromis species, but in none does the frequency 
of the variants exceed about 30 per cent. All 22 specimens of female H. chromogynos 
are " bicolor " . These fishes were collected from several different parts of the lake 
and include fishes at various stages of sexual development. 

Holotype. A female, 79 mm. standard length, from the Napoleon Gulf, near Jinja, 
Uganda. 

Description, based on 29 specimens (including the holotype) 50-110 mm. standard 
length. 

Depth of body 32-5-42-3 (M = 35-0) per cent of standard length ; length of head 
30-4-37-3 (M = 33-2) per cent. Dorsal head profile curved and moderately declivous. 
Preorbital depth 13-2-17-0 (M = 15-4) per cent of head length, least interorbital 
width 22-6-31-4 (M = 27-5) per cent. Snout slightly longer than broad, or less 
commonly, broader than long, its length 26-3-33-3 (M = 30-7) per cent of head ; 
eye 25-7-32-7 (M = 28-6) ; depth of cheek 17-9-24-6 (M = 21-6) per cent of head. 

Caudal peduncle 13-6-18-5 per cent of standard length, i-i-i-6 times as long as deep. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 213 

Mouth horizontal ; jaws equal anteriorly, the lower 1-1-1-5 (mode 1-3) times as 
long as broad, its length 30-0-34-4 (M = 32-5) per cent of head length. Lips thickened; 
posterior tip of the maxilla extending to the vertical through the anterior orbital 
margin or slightly beyond. 

Gill rakers short, 8 (less frequently 9 and rarely 6 or 7) on the lower limb of the 
anterior arch. 

Scales ctenoid ; lateral line with 31 (f.3), 32 (f.io), 33 (f.13) or 34 (f.3) scales. 
Cheek with 3 (less frequently 2 or 4) series. Six to 8 scales between the origin of 
the dorsal fin and the lateral line ; 8 or 9 between the pectoral and pelvic fin bases ; 
chest scales small. 

Fins. Dorsal with 24 (f.3), 25 (f.24) or 26 (f.2) rays, anal with n or 12, comprising 
XV-XVI, 8-10 and III, 8 or 9 spinous and branched rays for the fins respectively. 
Pectoral fin slightly shorter than the head ; first pelvic ray produced, variable in 
its posterior extension but usually reaching the spinous part of the anal fin. 

Teeth. In the outer row of both jaws, the teeth are slender and gently recurved. 
Fishes less than 65 mm. S.L., have only unequally bicuspid teeth ; individuals 
65-95 mm. S.L. have an admixture of bi- and unicuspid teeth in which either type 
may predominate. Fishes more than 95 mm. S.L. have only unicuspid teeth. There 
are 24-42 (modal range 30-32) outer teeth in the upper jaw. 

The inner teeth are tricuspid in fishes less than 95 mm. S.L. and unicuspid in 
larger individuals ; an admixture of both types may occur. These teeth are arranged 
in three rows (less frequently two or four) in both jaws. 

Lower pharyngeal bone triangular, the dentigerous area 1-1-1-4 (mode 1-2) times 
as broad as long. Occasionally the median series of teeth are enlarged and sub- 
molariform ; more frequently, only the posterior few pairs are markedly enlarged. 
In a few specimens, no median teeth are enlarged. 

Coloration. As mentioned above, H. chromogynos is unique in apparently having 
only " bicolor " females. The colour patterns of these fishes are variable, but are 
within the range known for other species with " bicolor " females. In preserved 
material, the yellowish-silver ground colour appears yellowish- white, silver or brown. 
The colours of live males are unknown. 

Coloration of preserved males. Ground colour greyish-brown to grey ; lips, lower 
jaw and the anterior part of the branchiostegal membrane, lighter ; six or seven 
faint transverse bars visible on the flanks and caudal peduncle ; a faint lachrymal 
stripe is often present. Dorsal fin with the spinous part dusky, lappets lighter ; 
soft part orange-yellow. Anal dusky on the basal half, orange-yellow distally, with 
one to three white ocelli arranged in a single row. Caudal fin dark, but with a broad, 
orange-yellow margin. Pelvic fins black on the outer half, orange mesially. 

Ecology : Habitat. H. chromogynos is probably confined to the littoral zone and 
to water less than 20 feet deep ; it has only been caught over a firm substrate (rock, 
sand or shingle). 

Food. One record of stomach and intestinal contents is available ; the main 
contents were the remains of Trichoptera larvae and sand-grain cases, but a few 
larval chironomids and baetids were also identified. 



214 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

Breeding. One female carrying embryos in the buccal cavity was recorded. Sexual 
maturity is reached at a standard length of 90-100 mm. in both sexes. 

Affinities. H. chromogynos may be related to H. chilotes (see p. 211). The similarity 
between these species is most pronounced when the non-lobed lip forms of H. chilotes 
are compared with H. chromogynos. Superficially, H. chromogynos resembles H. 
crassilabris (Blgr.) but the dentition of the two species is markedly different. 

The available specimens of H. paucidens Regan, from Lake Kivu, indicate a very 
close relationship between the two species ; the most marked difference is the shallower 
cheek of H. chromogynos (mean depth 21-6, cf. 27-6 per cent for H. paucidens). 
Unfortunately there is no information on the coloration of female H. paucidens 
or on the breeding colours of male fishes of either species. 

Study material and distribution records 

Museum and Reg. No. Locality Collector 

Uganda 
B.M. (N.H.).— 1958. 1. 16.83 . Napoleon Gulf, near J inj a . E.A.F.R.O. 

(Holotype) 

1958. 1. 16. 71-75 . Napoleon Gulf, 

near J inj a golf course 
„ 1958. 1. 16.76-81 . Napoleon Gulf, J inj a pier 

,, 1958. 1. 16. 82 . Napoleon Gulf, 

bay opposite J inj a pier 
1958. 1. 16. 84 . Entebbe Harbour 

1958. 1. 16.69 . Grant Bay 

,, 1958. 1. 16.67-68 . Ramafuta Is., Buvuma Channel . 

1906. 5. 30. 415-416 . Bugangu . Degen. 

1906. 5. 30. 407-41 2 . Bunjako 

Tanganyika 
1958. 1. 16.63-66 . Mwanza, Capri Bay . E.A.F.R.O. 

Kenya 

1958. 1. 16. 70 . Kisumu Harbour 

Haplochromis aelocephalus sp. now 

(Text-fig. 12) 

Holotype. A male 96 mm. S.L., from Igwe Island. 

An interesting feature of this species is its wide range of variation in head shape ; 
the more extreme individuals might well be considered distinct species were it not 
for the presence of annectent forms (Text-fig. 13). This variation is not correlated 
with sex or size. The most constant specific characters are the multiseriate dentition, 
the small scales on the pectoral region, and the thickened lips. 

Description. Based on the holotype and 21 other specimens, 63-120 mm. standard 
length. 

Depth of body 31-3-38-4 per cent of standard length ; length of head 33-0-38-6 
per cent. Dorsal head profile straight or very slightly concave, sloping gently ; physi- 
ognomy variable. Preorbital depth 14-7-19-4 (M = 17-2) per cent of head length, 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 215 

least interorbital width 21-9-26-4 (M = 24-8) per cent. Snout 1-1-1-2 times as long 
as broad, except in a few extreme individuals where it is 1-25-1-30 times as long as 
broad ; snout length 32-0-39-0 (M = 35-3) per cent of head. Diameter of eye shows 
fairly clear-cut negative allometry with standard length, 25-0-30-8 (M = 27-8) 
per cent of head in fishes 62-100 mm. S.L. and 23-2-25-7 (M = 24-5) per cent in 
larger individuals ; depth of cheek 17-3-24-7 (M == 20-5) per cent. 

Caudal peduncle 12-8-18-5 (M = 16-2) per cent of standard length, i-i-i-6 (modal 
range 1-4-1-5) times as long as deep. 

Mouth horizontal, lower jaw projecting slightly ; posterior tip of the maxilla 
not quite reaching the vertical through the anterior margin of the orbit, except in 
one specimen. Lips thickened and variable ; in a few fishes there are faint indications 
of a lobe-like swelling on the lower lip. In all specimens there is a pronounced sub- 




Fig. 12. Haplochromis aelocephalus ; holotype. Drawn by Miss D. Fitchew. 



mental thickening which extends posteriorly for a short distance. Lower jaw length 
apparently correlated with head shape, being greatest in the more extreme individuals; 
namely, in seven " extreme " specimens (Text-fig. 13) 42-5-48-5 (M = 45-1) per cent 
of head and in the remaining specimens 37-0-46-9 (M = 41-3) per cent. The length/ 
breadth ratio of the lower jaw 1-6-2-6 (modal range 1-8-2-0). 

Gill rakers short, 7-9 on the lower limb of the anterior arch. 

Scales ctenoid ; lateral line with 31 (f.i), 32 (f.5), 33 (f.io) or 34 (f.6) scales. Cheek 
with 3 or 4 (rarely 2) series. Six to 8 scales between the origin of the dorsal fin and the 
lateral line ; 8 or 9 (rarely 7 or 10) scales between the pectoral and pelvic fin bases ; 
chest scales small. 

Fins. Dorsal with 24 (f.3), 25 (f.17) or 26 (f.2) rays, anal with n (f.i), 12 (f.16) or 
13 (f.5), comprising XV or XVI, 9 or 10 and III, 8-10 spinous and branched rays for 
the fins respectively. Pelvic fins with the first ray produced. Pectoral fin shorter 
than the head. Caudal truncate or sub truncate. 

Teeth. The outer row in both jaws of fishes less than 65 mm. S.L. is composed of 
slender and slightly recurved bicuspid teeth ; specimens 65-95 mm. S.L. have an 



216 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

admixture of bi- and unicuspids in which either form may predominate, whilst in 
larger fishes, all the outer teeth are unicuspid. There are 24-42 (mode 32) teeth in 
the outer row of the upper jaw. The dental arcade in the lower jaw is narrow 
anteriorly ; in a few specimens it is rather acutely pointed and resembles that of 
lobe-lipped H. chilotes (see p. 209). 

Teeth in the inner series of both jaws are generally tricuspid in fishes less than 95 
mm. S. L. and unicuspid in larger individuals ; a mixture of both types is found in 
specimens of an intermediate size. There are three to five, rarely two (mode five), 
series of inner teeth anteriorly in the lower jaw and three to six (mode five) in the 
upper. The innermost series of the lower jaw is usually implanted so as to lie almost 
horizontally. 

Lower pharyngeal bone triangular, the dentigerous area 1-0-1-4 (mode 1-2) times 
as broad as long. Teeth in the median series are variable in form. In most specimens 




1 cm. 

Fig. 13. Haplochromis aelocephalus ; individual variant of head profile (extreme form). 

these teeth (especially the upper three or four pairs) are somewhat enlarged and weakly 
cuspidate ; the next most common variant has these teeth slightly enlarged and 
clearly cuspidate. Finally, in a few fishes the median teeth are unmodified and resem- 
ble the other teeth. Two exceptional fishes had the entire median series enlarged 
and molariform and the pharyngeal bone noticeably stouter. 

Coloration. The colours of living fishes are unknown. 

Preserved material : Sexually active males. Ground colour grey-black, chest and 
branchiostegal membrane black ; faint indications of a coppery sheen on the oper- 
culum and flanks. Dorsal fin black except for the light lappets and a colourless 
band outlining the soft part of the fin ; caudal black basally, light (? orange) distally ; 
anal fin dark on the basal half and light (? orange) distally, with three or four hyaline 
ocelli arranged in a single row. Pelvics black. Females, quiescent and juvenile males. 
Ground colour greyish-silver (in sexually quiescent males there is a faint trace of 
coppery sheen on the operculum and the chest is dusky) with, in some, an interrupted 
or continuous, dark, mid-lateral stripe and five or six transverse bars on the flanks. 
Dorsal and anal fins yellowish, slightly dusky on the proximal half in quiet males, 
but hyaline and faintly maculate in females and immature males. Pelvic fins yellowish 
or hyaline in females and immature males, dusky in quiet males. 



A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 217 



Distribution. Known only from Lake Victoria. 

Ecology : Habitat. The species has been found in relatively few localities and 
only in the Uganda waters of the lake. However, the available data suggest that 
H. aelocephalus is restricted to water less than 40 feet deep and to areas where the 
substrate is firm (sand and rock). 

Food. Sixteen of the 20 specimens examined had food in the stomach or intestines ; 
from these it would seem that H. aelocephalus preys on a variety of invertebrate 
animals, and possibly even small fishes. 

In ten specimens the predominant food organisms were insects (particularly 
dipterous larvae, but also Ephemeroptera [Povilla adusta] and Trichoptera larvae). 
The non-insect food identified was : in two fishes, oligochaet worms ; in one, the 
remains of a prawn (Caridina nilotica Roux) ; in another, fragments of plant-tissue and 
a few Ostracoda ; and in two others, numerous fragments of lamellibranch and gastro- 
pod shells. One exceptional individual contained the remains of a small cichlid fish. 

The presence of sand grains in the stomach and intestines of many individuals 
suggests that the species may be a bottom feeder. 

Breeding. No data are available. 

Affinities. Haplochromis aelocephalus shows no special affinity with any other 
Haplochromis species in Lake Victoria ; the less extreme individuals resemble 
members of the H. nigrescens species-complex of piscivorous-insectivorous predators. 
The multiseriate dentition, however, disqualifies H. aelocephalus from a place in this 
complex, but suggests relationship with species of the H. sauvagei-H. prodromus 
group, and particularly H. xenognathus. In shape, the teeth of H. aelocephalus 
are unlike those of H. xenognathus which have characteristically recurved tips 
(Greenwood, 1957). In certain cephalic characters, especially the shape of the lower 
jaw, the narrow lower dental arcade and the semi-lobate lips, H. aelocephalus 
approaches H. chilotes but in all other characters there is no obviously close relation- 
ship between the two species. 

Diagnosis. H. aelocephalus may be distinguished by the following combination 
of characters : proportions of the head ; a multiseriate dentition with the outer 
teeth slender and gently recurved ; lips somewhat thickened. 



Study material and distribution records 
Museum and Reg. No. 



Locality 



Uganda 



B.M. (N.H.).- 


1958. I . 16.244 


Igwe Isl. 


(Holotype) 




j 1 


1958. I. I6.2I5 


Ekunu Bay 


It 


1958. I . I6.2l6 


Entebbe, Bugonga Beach 


" 


1958. I. I6.2I7 


Beach near Nasu Point, 
Buvuma Channel 


,, 


1958. I . 16.218-224 


Igwe Isl. 


It 


1958. I . 16.225-228 


Bay opposite Jinja, 
Napoleon Gulf 


It 


1958. I. I6.232-233 . 


Pilkington Bay 


tt 


1958. I. 16. 234-235 . 


Buka Bay 


It 


1958. I. 16.236-242 


Napoleon Gulf, near Jinja 


,, 


1958. I. 16.243 


Unknown 



Collector 
E.A.F.R.O. 



218 A REVISION OF THE LAKE VICTORIA HAPLOCHROMIS SPECIES 

SUMMARY 

1. Seven species, which feed almost exclusively on the embryos and larvae of 
other Cichlidae, are discussed. Haplochromis obesus (Blgr.), H. maxillaris Trewavas, 
H. melanopterus Trewavas, H. parvidens (Blgr.) and H. microdon (Blgr.) are redes- 
cribed on the basis of new and more extensive collections. Two new species, H. cronus 
and H. cryptodon are described. 

2. Notes on the ecology and feeding habits of these species are given. 

3. The relationships of these species are discussed and it is concluded that the group 
has a polyphyletic origin. 

4. Four other species are considered. These are all insectivorous and do not 
appear to be closely related to the other species of Haplochromis in Lake Victoria. 
Haplochromis plagiodon Regan & Trewavas and H. chilotes (Blgr.) are redescribed, 
and two new species, H. chromogynos and H. aelocephalus are described. 

5. H. chromogynos is of particular interest since the normal female coloration is 
apparently the " bicolor " piebald which occurs as an infrequent and sex-limited 
mutant amongst the females of other and unrelated species. 

6. Both H. chilotes and H. aelocephalus are noteworthy for the wide range of 
individual variability which they show. 

ACKNOWLEDGMENTS 

I wish to acknowledge my gratitude and thanks to the Trustees of the British 
Museum (Natural History) for the facilities afforded me ; to the authorities of the 
Museum National d'Histoire naturelle, Paris and of the Museo Civico di Storia 
Naturale, Genoa, who graciously allowed me to study type-material in their collec- 
tions ; and to Mr. A. C. Wheeler of the Zoology Department, British Museum 
(Natural History) who was responsible for making several radiographs used in this 
study. I am especially indebted to Dr. Ethelwynn Trewavas for her most helpful 
advice and criticism. 

REFERENCES 

Graham, M. 1929. A Report on the Fishing Survey of Lake Victoria, 1927-1928, and Appen- 
dices. Crown Agents, London. 

Greenwood, P. H. 1954. O n two cichlid fishes from the Malagarazi River (Tanganyika) with 
notes on the pharyngeal apophysis in species of the Haplochromis group. Ann. Mag. nat. 
Hist. (12) 7 : 401. 

1956a. The monotypic genera of cichlid fishes in Lake Victoria. Bull. Br. Mus. nat. 

Hist., Zool. 3 : No. 7. 

1956&. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae). Part I. 

Ibid. 4 : No. 5. 

1957- A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae). Part II. 

Ibid. 5 : No. 4. 

Regan, C. T. 1922. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. 157. 

Regan, C. T. & Trewavas, E. 1928. Four new cichlid fishes from Lake Victoria. Ann. Mag. 

nat. Hist. (10) 2 : 224. 
Trewavas, E. 1928. Descriptions of five new cichlid fishes of the genus Haplochromis from 

Lake Victoria. Ibid. (10) 2 : 93. 




THE ROSAURA EXPEDITION 

1937-38 

CHAETOGNATHA 



JOHN S. COLMAN 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 8 

LONDON : 1959 



THE ROSAURA EXPEDITION 1937-38 
CHAETOGNATHA 



BY 



JOHN S. COLMAN 

Marine Biological Station (University of Liverpool) 
Port Erin, Isle of Man 



Pp. 219-253 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 8 

LONDON : 1959 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5 , No. 8 of the Zoological 
series. 




© Trustees of the British Museum, 1959 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued February, 1959 Price Twelve Shillings 



THE ROSAURA EXPEDITION 1937-38 
CHAETOGNATHA 

By JOHN S. COLMAN 



INTRODUCTION 

Genus SAGITTA Quoy & Gaimard, 1827 



Group 

(a) " Friderici " 

(b) " Bipunctata " 



(c) " Serratodentata 

(d) " Lyra " . 

(e) " Hexaptera " . 

(f) " Enflata" 

(g) " Planctonis " . 

(h) Ungrouped 



1. 

2. 

3- 

4- 

5- 
6. 

7- 
8. 

9- 
10. 
11. 
12. 

13. 

14. 

15- 
16. 







Number 






of 


Species 


Number 


hauls 


Sagitta friderici Ritter-Zahony, 191 1 


2 


2 


5. tenuis Conant, 1896 


177 


2 


5. bipunctata Quoy & Gaimard, 1827 


6 


5 


5. elegans Verrill, 1873 


42 


2 


S. robusta Doncaster, 1902 


14 


3 


5. serratodentata Krohn, 1853 


95 


4 


5. lyra Krohn, 1853 


no 


11 


S. maxima (Conant, 1896) 


167 


10 


S. hexaptera d'Orbigny, 1843 


299 


11 


5. decipiens Fowler, 1905 


29 


7 


5. enflata Grassi, 1881 


141 


13 


5. planctonis Steinhaus, 1896 


12 


2 


5. zetesios Fowler, 1905 


84 


10 


S. neglecta Aida, 1897 


7 


5 


5. pulchra Doncaster, 1902 


2 


1 


S. macrocephala Fowler, 1905 


107 


7 



Genus PTEROSAGITTA Costa, 1869 
17. Pterosagitta draco (Krohn, 1853) 



45 



Genus KROHNITTA Ritter-Zahony, 1910 

18. Krohnitta subtilis (Grassi, 1881) . n 

Genus EUKROHNIA Ritter-Zahony, 1909 

19. Eukrohnia hamata (Mobius, 1875) . 204 

20. E. fowleri Ritter-Zahony, 1909 . 277 



Total 



1831 



5 
9 

19 



This collection contains 1,831 individuals of 20 species (16 of Sagitta, 1 Pterosagitta, 
1 Krohnitta and 2 Eukrohnia) taken in 19 hauls. In the above list the order of 
species and the groups of Sagitta are based on Furnestin (1957 ) ; I have added the 



zool. 5, 8. 



n§ 



222 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

group " planctonis " (see David, 1956). The most distinct of these groups, such as 
" lyra " and " planctonis ", are undoubtedly of some value in making minor distinc- 
tions within the genus Sagitta, but no useful purpose would be served by raising 
them to subgeneric rank ; it would be only a matter of time before attempts would be 
made to raise such subgenera to generic status, and this would only create confusion. 

The full list of stations on the Rosaura expedition is given in Colman (1954). 

I am grateful to Dr. J. H. Fraser, Dr. E. L. Pierce and Mr. P. M. David for much 
helpful and interesting correspondence. Dr. Pierce and Mr. David both examined 
some of my specimens and sent me some of theirs, and Dr. Fraser kindly read through 
the typescript. I also wish to thank Dr. Ben Dawes for examining the parasites. 

Before the catches of chaetognaths are discussed in detail, the limitations of this 
collection should be pointed out. All the catches were made with open nets, so that 
in the case of the deep hauls there is no telling in most cases from what depth the 
specimens came. With one exception, all the deep hauls were made with a 2-metre 
stramin net ; the stramin mesh is too coarse for the reliable sampling of the smaller 
and more slender species, such as S. friderici, S. tenuis, S. bipunctata, S. robusta, S. 
serratodentata, S. decipiens, S. neglecta, S. pulchra, P. draco and K. subtilis, all of which 
were probably under-represented in the deep hauls. 

Each deep haul lasted about two hours, and each catch contained up to hundreds 
of fish and scores of molluscs, Crustacea etc., many of which were larger than the 
chaetognaths. Sorting was done single-handed ; all that I was able to do was to 
pick out specimens of about 30 different categories in the various phyla until I did 
not appear to be finding anything new. It is hoped that this procedure will give 
some idea of the relative abundance at each station of at any rate the larger chaetog- 
naths, but I do not think that I sampled adequately such smaller species as the stramin 
net was able to retain. 

Within the chaetognaths themselves the only sorting done on the spot was between 
red and colourless forms. All of Sagitta macrocephala and Eukrohnia fowleri were 
red, and there were occasional red individuals of S. enflata, S. zetesios and Pterosagitta 
draco. 

Owing to the length of the deep hauls, many of the specimens are more or less 
damaged ; this is much less true of the short hauls made near the surface with the 
small silk townet, which confirms the observation of David (1955, p. 240). In addition, 
all my samples (after killing with formalin) were preserved in 70% alcohol ; weak 
sea-water formalin should have been used, since alcohol encourages contraction and 
leads to damage. These factors have combined to make useless in many cases such 
taxonomically valuable structures as seminal vesicles, corona and fins, and to make 
many of my length-measurements of only relative value. Length-measurements in 
any case should be treated with some caution ; Russell (1932a, b ; 1933a, b) and 
Clarke et al. (1943) have shown that there is a very considerable difference in length 
between successive broods of Sagitta elegans and 5. setosa at different times of the 
year, and the same may well be true of other species which have not been studied 
with equal thoroughness. 

The hauls could not be planned much in advance, but were made when time 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 223 

and depth permitted. The depth of the net was calculated roughly from a visual 
estimation of the angle of the towing warp to the horizon, and cannot pretend to be 
accurate. It was my intention, at each haul, to send the net to a depth of about 
1,000 m., but several hauls had to be shallower than this ; sometimes there was not 
time to pay out and haul in enough wire, and sometimes there was no nearby sounding 
on the chart and the possible existence of sea-mounts had to be borne in mind. 

With the exception of the very young Sagitta elegans in Stations 4 and 7, every 
specimen has been examined for the following features : 

Overall length to the nearest o-i mm. (excluding tail fin) measured on a glass slide 
graduated in half-millimetres (some of the smallest were measured by eye to 0-05 mm.) . 

Length of tail, expressed as percentage of total length. 

Numbers of hooks, of anterior teeth and of posterior teeth ; whenever the two sides 
differed the larger number was taken, and I have included the smallest visible hook- 
rudiments. 

State of development of gonads. Several not dissimilar schemes exist for classifying 
the developmental stages of chaetognaths, notably those of Kramp (1917, 1939), 
Russell (1932), Thomson (1947), Faure (1953) and David (1955). The arbitrary 
division into stages of a continuous process is bound to be to some extent subjective 
so it is desirable for each author to define the stages which he uses. I have recorded 
the state of development of my specimens in six stages, as follows : 

Stage O : neither testes nor ovaries visible under X 100 magnification. 

I : testes visible, but no sperm-balls or spermatozoa free in tail coelom ; 
ovaries visible but very small. 
„ II : tail filled with spermatozoa, ovaries developing but eggs small. 
,, III : tail empty, ovaries further developed, with some eggs larger than others ; 

some eggs may even be full-sized. 
„ IV : tail empty ; all eggs full-sized ; in some cases, probably because of 
damage, many eggs are free in the trunk coelom. 
V : spent, the ovaries being reduced to crumpled remnants. 

Stage V seems to be almost confined to the genus Eukrohnia. This seems to imply 
that most, if not all, species of Sagitta die very soon after ovulation (or after the last 
ovulation in species such as S. enflata which are believed to go through more than 
one reproductive cycle in a lifetime). Indeed, van Oye (1931) states that this is so. 

The relative duration of the several stages, the speed of transition from one stage 
to the next, and the relative speed of development of testes and ovaries, vary not 
only between one species and another, but also to some extent between individuals 
of one species. I have occasionally found it convenient (especially in the case of 
Sagitta lyra, see below) to record two intermediate stages. 

Stages I— II : testes still retaining their shape, but some sperm-balls already free in 
tail coelom. 
" II— III : some spermatozoa still in tail, but ovaries quite as mature as in some 
specimens of stage III. 



224 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

DEEP-WATER HAULS 

Thirteen plankton hauls were made in deep water with 2-metre nets, 12 with the 
stramin net and one (Station 14) with a silk net of quarter-inch mesh. The latter net, 
as might be expected, caught very few chaetognaths, and one of the stramin net 
hauls (Station 41) was spoilt by containing a large shark. The details of the samples 
from these stramin net hauls are shown in Table I. 

Station 8 was in near-Arctic waters south-west of Greenland, Station 13 was in 
temperate waters between the Gulf Stream and the continental shelf off the eastern 
United States, and the remainder were in tropical waters ; Stations 15-33 wer e 
in the Caribbean Sea, and Stations 42-46 in the central Atlantic. Station 8 took place 
at night ; all the others were in daylight. 

The sample from Station 8 can be compared with the catches from the Godthaab 
Expedition of 1928, described by Kramp (1939). In deep water south-west of Cape 
Farewell Kramp found that Sagitta maxima and Eukrohnia hamata were abundant, 
with " 5. planctonis " (= S. zetesios) (see David, 1956) and E. fowleri less common. 
In Rosaura Station 8 the same four species occurred in the same order of abundance, 
but to them can now be added the red, deep-water species, S. macrocephala. It is 
surprising that Kramp found none of the latter species, for the Godthaab's hauls 
were made at all depths down to 2,000 m., a good deal deeper than mine which I 
estimated to reach 1,300 m. 

A submarine ridge joins southern Greenland to southern Baffin Land about 65 ° N. 
According to Kramp neither "Sagitta planctonis" (= S. zetesios) nor Eukrohnia 
fowleri is found north of the ridge. S. maxima and E. hamata, however, both extend 
northwards as far as Ellesmere Land and North Greenland, and in these northern 
waters they are joined by S. elegans arctica as an offshore species. Further south in 
the Davis Straits the latter species is confined to inshore waters ; I found no 5. 
elegans in Station 8, but did catch some very young juveniles (Stage O) in Stations 4 
and 7 near the Greenland coast. 

Station 13 was taken about 60 miles north-east of Cape Hatteras, and was thus 
only just outside the area covered by Pierce (1953). To judge by the surface tempera- 
ture (17*8° C.) and the fact that the ship was being set to the southward, Station 13 
must have been in the " slope " water between the Gulf Stream and the continental 
shelf, and the position would correspond roughly to Pierce's " Outer Shelf Zone ". 
Pierce recorded 11 species from this zone, and I identified eight species from the 
sample in Station 13, but only three species are common to the two lists, namely 
Sagitta enflata, 5. lyra and 5. bipunctata, as is shown on p. 225. 

The five species in the Rosaura sample which were missing from Pierce's list all 
inhabit moderately deep or deep water ; few of Pierce's samples came from deeper 
than 200 m. Of the eight species in Pierce's list which were not found in Station 
13, seven are small species which would not be caught in large numbers by a stramin 
net (though most of them did appear from time to time in other Rosaura samples). 
The remaining one, S. hexaptera, is a large species which throughout the Caribbean 
and the central Atlantic was one of the most regularly occurring in the Rosaura 
samples, and was often the most numerous. Pierce records it as " rare " in the Cape 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



225 





Pierce (1953) 


Rosaura 




Outer Shelf Zone 


Station 13 




( + + +, common 


(Numbers 




+ +, occasional 


in 


Species 


+ , rare) 


sample) 


Sagitta minima 


+ + + 




S. serratodentata 


+ + + 






S. helenae 


+ + + 






S. tenuis 


+ + 






S. hexaptera . 


+ 






Krohnitta pacifica . 


+ + + 






K. subtilis 


+ 






Pterosagitta draco . 


+ + + 






Sagitta enflata 


+ + + 




H 


S. lyra . 


+ 


2 


S. bipunctata 


+ + + 


1 


S. maxima 




12 


S. zetesios 




5 


S. macrocephala 




4 


Eukrohnia hamata . 




5i 


E. fowleri 




] 


C4 



Hatteras-Cape Fear region, and it has been occasionally recorded from as far north 
as Nova Scotia (Pierce quotes Huntsman, 1919) ; its absence from Rosaura Station 
13 may indicate that it is common only to the southward of Cape Hatter as. 

At Station 8 the surface temperature was 4-8° C, and at Station 13, 17-8° C. All 
the other offshore hauls with the stramin net were made in tropical waters with the 
surface temperatures ranging from 25-5 to 28-3° C, and throughout the extensive 
area from Central America to the eastern central Atlantic the catches on the whole 
resembled one another closely. 

Of the species large enough not to be able to escape through a stramin net, Sagitta 
hexaptera, S. enflata, S. maxima, S. lyra and 5. zetesios occurred regularly in all or 
nearly all of the samples, and of these 5. hexaptera was usually the most abundant. 
S. maxima, though usually present, seems to have been distinctly scarcer in tropical 
waters than in the Davis Straits (Station 8) . 

Another sizeable species, 5. planctonis, was found only in the samples from the two 
easternmost stations (45 and 46) between St. Paul's Rocks and the west coast of 
Africa, as was the smaller and more slender 5. friderici. Eukrohnia hamata, which 
dominated the catches in the two most northerly stations (8 and 13), was scarce in 
the tropics, providing only 14 specimens in three out of nine hauls. 

Of the smaller species, Sagitta decipiens and Pterosagitta draco occurred with 
surprising regularity and must have been abundant everywhere. The paucity of the 
other species does not necessarily indicate that they were rare ; a stramin net would 
let most of them through, and my sampling technique was not adequate for such as 
were caught. 

A special mention may be made of the two red species, Sagitta macrocephala 
and Eukrohnia fowleri. Of the nine tropical hauls, the estimated depth of the 
net was at least 1,000 m. in five of them, but not more than 900 m. in the other 



226 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

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THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 227 

four. S. macrocephala occurred in none of the latter, but provided 82 specimens in 
four of the deeper hauls ; E. fowleri was represented by two specimens in one 
(900 m.) of the less deep hauls, but by 253 in all five of the deeper ones. This fairly 
sharp division between 900 m. and 1,000 m. may show that my estimates of net-depth 
are at any rate reasonably self-consistent. 

INSHORE HAULS 

Six short hauls were made near the surface with a small silk townet of about 60 
meshes to the inch. The details of the samples from these are shown in Table II. 

The first two of these hauls (Stations 4 and 7) were taken in Greenland coastal 
waters. The first was, it is true, over fairly deep water (550 m.), but the coast was 
less than 10 miles away, and the catch looked like inshore plankton, being dominated 
by medusae and ophioplutei. At Station 7 there were few plutei, but again medusae 
were conspicuous. The only chaetognaths in these catches were some very young 
(stage O) Sagitta elegans, presumably the subspecies arctica ; since this species did 
not appear in the offshore sample at Station 8, these observations tend to confirm 
Kramp's (1939) statement that in the Davis Straits 5. elegans is confined to coastal 
waters. 

The catch at Station 23 in Belize Harbour (British Honduras) calls for little com- 
ment. It was made over shallow water in bright sunshine, and the few chaetognaths 
caught (one S. neglecta and six 5. enflata) were all very young ; the largest of the 
S. enflata was only 3-1 mm. long, and the others were post-larvae still with relatively 
very large ventral ganglia. The rest of the catch was typical of inshore waters, and 
was dominated by medusae, post-larval polychaetes, and the larvae of decapods 
and molluscs. 

The next two hauls (Stations 38 A and 40) are of considerable interest. They were 
both taken in the complex system of rivers and creeks in British Guiana where the 
water is very opaque (more so than the most turbid water in the Tamar Estuary, 
for example), the salinity low (10-9 and 13-9/00), and the plankton abundant, varied, 
and entirely marine in character. The most numerous groups in the plankton were 
young fish and fish larvae, penaeid prawns, copepods and chaetognaths, all of the 
last group being Sagitta tenuis. These tidal waterways are discussed at greater 
length by Colman & Cooper (1954), and 5. tenuis will be considered below in the sec- 
tion devoted to the individual species. 

The last silk net haul to contain any chaetognaths, Station 44, was made a few 
feet off St. Paul's Rocks, the tiny, isolated land-mass in the middle of the " waist " of 
the Atlantic. The shelf from which the rocks project is only a few hundred metres 
across and it supports a generous growth of algae, hydroids, polyzoa, sponges etc., 
accompanied by other invertebrates. At the time of the Rosauras visit the equatorial 
current was sweeping between the rocks from east to west at about 2 knots, but the 
plankton seemed to contain a local concentration of neritic forms, including mysids, 
orange, blue and variegated copepods, and post-larval gastropods. The five species 
of chaetognath, however, were all widely-distributed oceanic species ; in the sample 
of in there were a single Sagitta bipunctata, several each of 5. hexaptera, S. enflata 
and Pterosagitta draco, and 90 5. serratodentata. 



228 THE ROSA URA, EXPEDITION 1937-38: CHAETOGNATHA 

This population is not unlike that in the surface waters at Fernando de Noronha, 
some 300 miles (500 kilometres) south-west of St. Pauls' Rocks. At Fernando de 
Noronha in 1954 (Hosoe, 1956) Sagitta serratodentata atlantica made up two-thirds of 
the chaetognaths, and was followed in order of abundance by S. enflata, Pterosagitta 
draco, Krohnitta pacifica and Sagitta hexaptera. 

NOTES ON THE SEVERAL SPECIES 

The reference under each species heading are to good descriptions which I have 
found useful in identification ; they are not lists of synonymies. The latter will be 
found in Ritter-Zahony, 191 1 ; Michael, 191 1, 1919 ; Johnston & Taylor, 1921 ; 
Burfield & Harvey, 1926 ; Burfield, 1930, 1950 ; Kramp, 1939 ; Tokioka, 1939, 
1940a, 19406 ; Thomson, 1947 ; Pierce, 195 1 ; Vannucci & Hosoe, 1952 ; David, 
1955, 1956, 1958 ; Furnestin, 1957. 

Whenever a species has been sufficiently numerous, I have shown the details of 
head armature etc. separately for young forms (stages O and I) and adults (stages II 
and up) . This seems to me to be preferable to the more usual method of showing the 
range of variation within different size-groups (see, for example, Thomson, 1947), 
since the development of head armature is more a function of maturity than of 
size. Furthermore, I have shown, in addition to the range of variation, the numerical 
distribution within the range ; this method can show up specific and racial differences 
better than can the range of variation alone. At the end of the paper I have summa- 
rized the head armatures and tail lengths of the adults of the species in the Rosaura 
collection. 

Sagitta friderici Ritter-Zahony 

Ritter-Zahony, 191 1 : 19. 

Fraser, 1952 : 14. 

Vannucci & Hosoe, 1952 : 14. 

Faure, 1953 : 26. 

Furnestin, 1954 ; 1957 : 113. 

Mme. Furnestin (Faure, 1953 and Furnestin, 1954) has shown that 5. friderici 
is distinguishable from S. bipunctata, but there has been some question as to whether 
S. friderici is separable from 5. tenuis ; the latter point will be discussed in the section 
below on S. tenuis. Only two specimens of 5. friderici occurred in the Rosaura 
collection, one from each of Stations 45 and 46. Both these were over deep water in 
the eastern central Atlantic, and bear out the suggestion of Vannucci & Hosoe 
(1952) that this species is not so neritic as was suggested by Faure (1953) . 

Length Tail 

(mm.) (% length) Hooks 

12 "j 26 . 8 

6-8 . 25 7 

The best character by which this species can be separated from S. bipunctata 
seems to be the propinquity of the seminal vesicle to the posterior fin ; in S. bipunctata 
there is a gap, but in S. friderici there is not. 



Ant. 


Post. 




teeth 


teeth 


Stage 


8 


18 


II 


6 


12 


I 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 229 

Sagitta tenuis Conant 

Michael, 191 1 : 72. 
Pierce, 195 1 : 219. 
Tokioka, 1955 : 57. 

This is not yet a well-known species, and several authors have cast doubt on its 
validity ; the only sound work on its distribution has been that of Pierce (1951, 
*953> 1958) and of Bieri (1957). 

Conant (1896) described S. tenuis rather inadequately from specimens taken in 
Kingston Harbour, Jamaica. Ritter-Zahony at first (19106) regarded it as a synonym 
for 5. bipunctata, but later (191 1) listed it among a number of doubtful species. 
Michael (191 1) re-examined Conant's material and kept S. tenuis separate from 
S. bipunctata. Pierce (195 1) redescribed 5. tenuis, but tentatively regarded it as a 
synonym for S. friderici on the grounds of its resemblance to Ritter-Zahony's 
(191 1) description of the latter. Fraser (1952), however, in his valuable discussion 
of the confusion between 5. bipunctata and other species, feels that 5. tenuis should 
be kept separate from S. friderici if only because of the great difference in size between 
the two : S. tenuis less than 8 mm., mature S. friderici more than 10 mm. Tokioka 
(1955), after expression some preliminary doubt, comes to the conclusion that S. 
tenuis and S. friderici are both valid species, while Bieri (1957) suggests that, in 
the coastal waters of Peru and Lower California at any rate, the two species represent 
the ends of an unbroken sequence of ecotypic variation. Finally, Furnestin (1957) 
firmly separates the two, placing them both, together with 5. setosa Miiller, S. hispida 
Conant and 5. helenae R.-Z., in her " groupe friderici " of neritic species. 

After examining 103 specimens from Station 38A and 73 from Station 40 (both 
from tidal rivers in British Guiana) I am in no doubt that my material is identical 
with Pierce's S. tenuis, and that S. tenuis is a valid species separate from 5. friderici. 

From Station 38A there were 103 specimens, including 7 stage I, 48 stage II and 
48 stage III ; in the 73 specimens from Station 40 there were no stage I, 22 stage II 
and 51 stage III. In the accompanying tables I have omitted those of stage I, 
because they had not yet acquired their full complement of teeth ; the two lots are 
shown separately because there are certain small differences between them, viz. 

(1) Front teeth : most of those from Station 38A have 5, most of those from Station 
40 have 6. 

(2) Anterior fins : these begin somewhat further forward in station 38 A than in 
Station 40, and are somewhat longer ; they end at about the same level in both lots. 

In a separate table I also show the anatomical details of six 5. tenuis kindly sent 
to me by Dr. Pierce, collected by him at Cedar Keys, Florida. As regards length, 
numbers of back teeth and front teeth, position of hind-end of anterior fins, and length 
of posterior fins, there is no difference between those from Florida and those from 
British Guiana. On the other hand the Floridan specimens have somewhat shorter 
tails (26-29% v. 27-33%), fewer hooks (7, 8 v. 8, 9), shorter anterior fins (16-21% 
v. 18-26%) which also begin further back (34-38% v. 29-36%), and less of the pos- 
terior fin on the tail (52-65% v. 58-75%). These differences do not seem more signi- 
ficant, however, than those already noticed between the two lots from British 

ZOOL., 5 8. II§§ 



230 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 













Sagitta tenuis 




















Tail 






Anterior 


Posterior 




Length 






A 


Hooks 




teeth 




teeth 




r 


^4 




X 






/o 




x 


^ 


A 




X 


f 






■\ 


r 


r > 


r 


'x 


mm. 


38A 


40 


Total 


lgth. 38A 40 Total 


No. 38A 40 Total 


38A 40 Total 


38A 


40 Total 


>7 


I 


— 


1 


<2y 


5 1 6 


3 — — - 










6-5-7 


I 


— 


1 


27 


9 1 10 


4 — — - 




10 — 10 


— 


— — 


6-65 


3 


2 


5 


28 10 7 17 


5 — — - 




69 13 82 


— 


— — 


5-5-6 


10 


6 


16 


29 1 


3 14 27 


6 — — - 




14 40 54 


2 


— 2 


5-5'5 


18 


14 


32 


30 21 7 28 


7 1 — 


1 


— 17 17 


3 


— 3 


4-5-5 


27 


36 


63 


31 19 14 33 


8 55 44 99 


— 1 1 


6 


1 7 


4-4-5 


25 


14 


39 


32 10 18 28 


9 38 28 66 


— — — 


6 


3 9 


3-5-4 


11 


1 


12 


33 

34 

>34 


4 9 13 
415 
1 1 2 


JO — — - 

11 — — - 

12 — — - 

13 — — - 

14 — — - 

15 — — ~ 

16 — — - 






14 
18 
20 
13 
17 
1 

4 


10 24 

12 30 

20 40 

12 25 

7 24 

6 7 

1 5 










Anterior fins 

X 






1 

Gap betwe 




Anterior end 


Posterior end 


Length of fin 


en fins 




x 








A. 


x 






X 




r 

/o 






*\ 


t 

/o 


^ 


i 

/o 




*\ ( 

/o 




1 


lgth. 38A 


40 Total 


lgth. 38A 


40 Total 


lgth. 38A 40 


Total lgth. 


38A 


40 Total 


28 


4 


— 


4 


49 1 


2 3 


16 — 1 


1 


L • 3 


I 


— 1 


2 9 


8 


1 


9 


50 4 


1 5 


17 — 4 


4 


I • 4 


7 


9 16 


30 


10 


2 


12 


5i 5 


5 10 


18 4 4 


t 


5 • 5 


22 


10 32 


31 


11 


6 


17 


52 18 


13 31 


19 12 13 


21 


> . 6 


12 


16 28 


32 


23 


16 


39 


53 7 


15 22 


20 9 19 


2* 


J • 7 


15 


15 30 


33 


18 


15 


33 


54 18 


14 32 


2J IO 16 


2( 


> . S 


13 


11 24 


34 


6 


17 


23 


55 8 


10 18 


22 19 6 


22 


\ . 9 


12 


5 17 


35 


6 


9 


15 


56 18 


7 25 


23 14 6 


2( 


> . IO 


8 


5 13 


36 


2 


5 


7 


57 5 


4 9 


24 7 3 


1( 


) . 11 


3 


1 4 


37 


1 


2 


3 


5<? 3 


2 5 


25 8 — 


\ 


J . 12 


— 


— — 


38 


2 


— 


2 


59 1 


— 1 


26 7 1 


i 


J . *3 


— 


I 1 


39 


2 




2 


60 2 
>6o 3 


— 2 

— 3 


27 1 — 

2<S 2 — 


1 


L 
5 














Posterior fins 






















x 






^ Percenl 


"a Or#» 


r 
















,<ige 


Anterior end 


Posterior end 


Length of fin 


of fin on 


tail 




x 






x 


JL 






X. 




% 






> 


% 


1 


% 




% 




■ ** 


lgth. 3 8a 


40 Total 


lgth. 38A 


40 Total 


lgth. 38A 40 


To1 


:al lgth. 


38A 


40 Total 


56 


— 


1 


1 


84 2 


1 3 


21 1 — 


] 


1 . <5* 


5 


2 7 


57 


9 


6 


15 


<*5 5 


3 8 


22 3 1 


4 


t • 5S-59 


4 


3 7 


58 


4 


8 


12 


<S6 7 


13 20 


23 3 4 


» 


7 . 60-61 


5 


2 7 


59 


8 


10 


18 


<?7 19 


21 40 


24 5 2 


p 


7 . 62-63 


12 


8 20 


60 


19 


17 


36 


** 32 


23 55 


25 11 6 


i: 


7 . 64-65 


12 


11 23 


61 


15 


10 


25 


89 12 


9 21 


26 15 12 


2: 


7 . 66-67 


20 


14 34 


62 


15 


10 


25 


90 13 


2 15 


27 25 20 


41 


> . 68-69 


13 


15 28 


63 


7 


4 


11 


9* 3 


1 4 


2# 12 13 


21 


> . 70-71 


12 


8 20 


64 


7 


2 


9 


92 1 


— 1 


29 10 9 


V 


> • 72-73 


3 


3 6 


65 


7 


2 


9 


93 1 


— 1 


30 8 5 


U 


^ . 74-75 


3 


3 6 


66 


2 


3 


5 






3-f 1 1 




8 . 76-77 


2 


2 4 


67 


— 


— 


— 






32 1 — 


] 


I . >77 


4 


2 6 


68 


1 


— 


1 
















69 


1 


— 


1 


% 















THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



231 



Six Sagitta tenuis from Cedar Keys, Florida (Collected by Dr. Pierce, 15.^.49) 





Tail 




Ant. 


Post. 




Length 


(% 


Hooks 


teeth 


teeth 




(mm.) 


lgth.) 


No. 


No. 


No. 


Stage 


6-05 


26 


8 


6 


15 


III 


6-05 


26 


8 


5 


12 


III . 


5-95 


28 


7 


6 


14 


Ill . 


5*5o 


29 


7 


5 


11 


Ill . 


5-50 


27 


8 


5 


13 


Ill 


5-35 


29 


8 


6 


13 


III . 





Ant. 






Post. 


per- 
centage 


Ant. fin. 


fin. 


Gap 




fin. 


of post. 


(% 


(% 


(% 


Post. fin. 


(% 


fin on 


lgth.) 


lgth.) 


lgth.) 


% lgth.) 


lgth.) 


tail 


37-54 


17 


9 


63-88 


• 25 


• 56 


38-55 


17 


6 


61-88 


. 27 


52 


36-54 


18 


9 


63-89 


. 26 


• 65 


35-51 


16 


8 


59-86 


. 27 


• 56 


34-55 


21 


7 


62-87 


• 25 


• 56 


35-53 


18 


7 


60-86 


. 26 


■ 58 



Guiana ; in each instance there is an overlap, and there seem to be no grounds for 
regarding the Floridan and British Guianan specimens as belonging to different 
species. As mentioned above, Faure (1953) gives good reasons for keeping 5. tenuis 
separate from 5. friderici, but she also casts some doubt on the identity of Pierce's 
with Conant's 5. tenuis. Pierce (1951), however, compared Conant's Jamaican 
syntypes directly with his own Floridan material and this, he states, " revealed that 
they were the same species ". 

Hitherto not very much has been known about the distribution of 5. tenuis. 
Pierce (195 1) found it abundant on the west coast of Florida, where it flourished in 
a salinity of 35 % but tended to avoid the most inshore waters where the salinity 
fell below about 25 % . (In the latter positions it was replaced by another small 
species 5. hispida Conant.) Pierce (1953, 1958) also described its occurrence off 
North Carolina. Here 5. tenuis is markedly neritic ; its seaward distribution is 
limited by the 36 % isohaline, and it is most abundant in such places as the southern 
mouth of the Pamlico Sound, where it occurs in salinities as low as, but not lower 
than, 22*7% . The type locality of S. tenuis is Kingston Harbour, Jamaica, and other 
places mentioned by Pierce are various inlets and estuaries in the south-eastern 
United States where the salinity will be reduced, but not very reduced. Bieri 
(1957) records 5. tenuis in the coastal current of Peru in salinities of 32-34 % and 
temperatures of 18-20 C. 

In the areas worked by Pierce and Bieri, then, S. tenuis seems to be confined to the 
salinity range 23-36 % . The Rosaura specimens show that in British Guiana it 
flourishes and breeds in salinities as low as io% . It would clearly be desirable to 
have a series of samples of S. tenuis, with salinity measurements, from various inlets 
and estuaries round the coast of Central America and from islands other than Jamaica. 



Sagitta bipunctata Quoy & Gaimard 



Ritter-Z&hony, 191 1 : 16. 
Michael, 191 1 : 41. 
Ghirardelli, 1950: 115. 
Fraser, 1952 : 13. 
Faure, 1953 : 36. 
Furnestin, 1957 : I 7 I - 



232 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



5. bipunctata has often been confused with other species (see the valuable discussion 
on this point by Fraser (1952)), but Mme. Furnestin (Faure, 1953 ; Furnestin, 1954, 
1957) has clearly established its identity and diagnostic limitations. 

There were only six specimens in the Rosaura collection (it is too slender a species 
for the stramin net) ; one was from Station 13 off the eastern United States, and the 
others were all caught in the central Atlantic. Their specifications were as follows : 



Length 



Tail 



Ant. 



Post. 



mm.) 


(% length) 


Hooks 


teeth 


teeth 


Stage 


n-8 


25 


8 


4 


10 


II 


hi 


27 


10 


7 


15 


III 


11 -o 


25 


8 


7 


13 


II 


io-8 


24 


9 


6 


16 


II 


9-6 


28 


9 


6 


16 


II 


6-7 


24 


9 


7 


14 


III 



The smallest specimen had already reached stage III in maturity. 



Sagitta elegans Verrill 
Ritter-Zahony, 191 1 : 14. 
Fraser, 1952 : 7. 
Fraser, 1957. 

As mentioned earlier (p. 227) 43 young specimens were taken close to the Greenland 
coast. The largest was 9 mm. long, all were stage O juveniles, and none had acquired 
the adult number of teeth. 

Sagitta robusta Doncaster 
Ritter-Zahony, 191 1 : 16. 
Michael, 1919 : 259 (as S. ferox). 
Burn eld & Harvey, 1926 : 100. 
Tokioka, 1939 : 127 (as 5. ai). 
Tokioka, 1940a : 4 (as 5. ai). 
Thomson, 1947 : 13. 

" The identity of S. robusta is most confused " (Thomson, 1947). This statement 
is true if S. ferox Doncaster is regarded as a species separable from S. robusta, a 
point about which there has been much argument (see below) ; otherwise S. robusta 
is an easily recognized species, with its unusually large head and its stiff, slim body. 

Doncaster (1902), when he described both species from the Indian Ocean, himself 
suggested that they might prove to be identical. Ritter-Zahony (191 1) united them 
under the name S. robusta. Burfield & Harvey (1926), after re-examining Doncaster's 
original material, concluded that on balance it was better to regard 5. ferox as a 
synonym of 5. robusta ; they saw a large number of specimens. 

On the other hand Fowler (1906), and after him Michael (191 1), described a Far 
Eastern species as 5. ferox and regarded it as separable, on the evidence of published 
descriptions, from 5. robusta ; according to Thomson (1947), however, the 5. ferox 
of Fowler and of Michael was really the 5. robusta of Doncaster. Tokioka (1939 
and 1940a), working in Japanese waters, found two species which he named 
5. robusta and S. ai, but according to Thomson Tokioka's robusta is really Doncaster's 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



233 



ferox, and ai is Doncaster's robusta. Finally Thomson says that in the waters south- 
east of Australia " the two are easily differentiated ". 

The head armatures of 5. robusta and 5. ferox are almost identical, but Thomson 
(1947) gives the following characters on which to separate the two species (all 
figures expressed as percentages of total length) : 







Width 


Length of 


Length of 




Width 


of head 


ant. fin 


Post, fin 


5. robusta . 


6-1-6-6 


9-4-11-4 


25-5-30-4 


25-4-30-8 


S. ferox 


5-4-5'S 


7-7-8-3 


21-1-22-7 


25-0-27-0 



Tokioka (1939) stresses another character, namely that the mature seminal vesical 
has a large head in his 5. robusta (= 5. ferox Doncaster) but not in his S. ai (= S. 
robusta Doncaster). 

Unfortunately the 14 specimens of 5. robusta from the Rosaura collection seem 
only to confuse the issue ; some of their characters are those of Thomson's 5. robusta, 
but others of his S. ferox. 

Width. After their long stay in alcohol the Rosaura specimens are bilaterally 
concave, and no longer suitable for width-measurement. 

Length. They show no obvious signs of longitudinal contraction. The largest 
Rosaura specimen (stage II) measured 11-5 mm.; this suggests S. ferox. (Doncaster's 
ferox reached 13 mm., but his robusta 16 mm., while Thomson records a robusta 
as long as 22 mm.) 

Width of head. This ranged from 9-5 to 12-2% of the body length, and 10 of the 
14 specimens lay within the range io-o-ii-i%. This clearly suggests robusta. 

Length of anterior fins. The anterior fins could be measured in 12 specimens, and 
ranged from 17-0 to 20-0% ; this as strongly suggests ferox. 

Seminal vesicles. Only three specimens had vesicles which seemed to be mature, 
but none of them had a large head. As far as it goes this suggests robusta. 

It should be noticed that each of these features in the Rosaura specimens belongs 
definitely to one or other of Thomson's species ; they are not intergrades between 
the two. 

The other anatomical details of the Rosaura specimens are as follows : 



^ength 


Tail 




Ant. 


Post. 




(mm.) 


(% length) 


Hooks 


teeth 


teeth 


Stage 


n-5 


25 


8 


10 


12 


II 


II-2 


26 


7 


10 


12 


II 


10-9 


26 


7 


8 


12 


II 


io-6 


26 


7 


9 


12 


II 


10-4 


24 


7 


8 


12 


I 


10-4 


27 


8 


9 


12 


I 


io-o 


26 


7 


8 


15 


II 


9-6 


26 


7 


8 


11 


II 


9-5 


26 


7 


7 


12 


I 


9-i 


24 


8 


8 


13 


I 


8-6 


28 


8 


9 


13 


I 


8-4 


27 


7 


7 


12 


I 


8-3 


27 


7 


7 


12 


I 


5.8 


28 


9 


7 


11 


O 



234 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

Now, those who have found it difficult or impossible to separate S. ferox from 
5. robusta (Doncaster, Ritter-Zahony, Burfield & Harvey, Colman) have worked on 
material from either the Indian or the Atlantic Ocean. The Pacific Ocean, on the 
other hand, has provided the material for those (Fowler, Michael, Tokioka, Thomson) 
who have distinguished these two species. This discrepancy could, perhaps, be 
explained on the supposition that S. robusta is a single, though somewhat variable, 
species in the Indian and Atlantic Oceans, but is evolving in the Pacific Ocean into 
two recognizable subspecies, S. robusta robusta and S. robusta ferox. Such an explana- 
tion, if true, would be surprising, since the Indian Ocean is, in general, more closely 
linked faunistically with the Pacific than with the Atlantic. 

Sagitta serratodentata Krohn 
Ritter-Zahony, 191 1 : 22. 
Michael, 191 1 : 39. 
Thomson, 1947 : I 5- 
Ghirardelli, 1950 : 120. 
Fraser, 1952 : 8. 
Fraser, 1957. 
Furnestin, 1957 •' J 47- 

5. serratodentata was for long regarded as a single variable species. Tokioka 
(19406) recognized two forms or subspecies 5. s. atlantica and 5. s. pacifica in 
Australian waters, and Thomson (1947) added a third form S. s. tasmanica from 
the same region. Fraser (1952) states that all three varieties occur together in Scottish 
waters, and doubts their subspecific validity ; but later (1957) he ascribes the variety 
tasmanica, in the eastern Atlantic, to more northern waters, and atlantica to warmer, 
southern and more saline waters. Furnestin (1957) firmly divides the species into 
three separate ones : S. serratodentata (= /. atlantica), S. pacifica and 5. tasmanica. 

The main character on which this separation is made is the structure and appearance 
of the mature seminal vesicle. There were 31 Rosaura specimens with mature seminal 
vesicles, and these were all of the atlantica type ; so fortunately, whatever the final 
decision on these taxa maybe, the Rosaura material will still belong to S. serratodentata, 
the type-locality of which is Messina. 

There were 95 Rosaura specimens, of which only five were caught by the stramin 
net (at Station 32 in the western Caribbean, and at 45 and 46 in the eastern central 
Atlantic) ; the other 90 all came from one short haul with the small silk net within 
a few yards of St. Paul's Rocks just before noon in bright sunshine. Table II (p. 226) 
shows that at the time of the Rosaura s visit 5. serratodentata was by far the commonest 
surface-living chaetognath at St. Paul's Rocks. 

The smallest specimens measured 4-4 mm. (stage O), and the largest 9-6 mm. 
(stage III). The size-range and frequency of occurrence of the maturity stages were 
as follows : 

Size-range 

Stage (mm.) Number 

O . 4-4-6-9 . 5 



I 

II 
III 
IV 



5-8-8-5 
6-8-9-5 
7-3-9-6 

7-8 5 



30 
41 
18 

1 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 235 

Other details were as follows : 

Sagitta serratodentata 





Length 
_ ._■*■ 




0/ 
/o 


Tail 
(as % of length) 

O, I II-IV Total 




Hooks 

A, 




Anterior teeth 




Posterior teeth 


mm 


O.I 


II-IV Total 


No. 


0,1 


II-IV Total 


0,1 


II-IV 


Total 


No. 


0,1 


II 


-IV Total 


r* 9 

8-9 
7-8 
6-7 


5 
8 


13 

34 

12 

1 


13 

39 

29 

9 


19 

20 

21 
22 


1 
2 

4 


7 


1 . 

2 . 

11 ! 


5 
6 

7 
8 


4 

27 

4 


28 

30 

2 


32 

57 

6 


2 
2 
8 
11 


1 
17 


2 

2 

9 

28 


5-J2 

14 
15 


5 
7 
3 
7 




— 5 

— 7 
2 5 
8 15 


5-6 


3 


— 


3 


23 


3 


11 


14 . 


9 


— 


— 


— 


10 


33 


43 


. 16 


10 




10 20 


4S 


2 


— 


2 


24 


11 


18 


29 . 


10 


— 


— 


— 


2 


8 


10 


■ 17 


1 




8 9 










25 
26 


4 
9 


21 
3 


25 . 
12 . 


11 
12 














1 


1 


. 18 
• ^9 


2 




10 12 
6 6 










27 


— 


— 


— 
















20 


— 




6 6 










28 


— 


— 


— 
















21 


— 




2 2 










29 


1 


— 


1 
















22 
25 


— 




7 7 
1 1 



Sagitta lyra Krohn 
Ritter-Z&hony, 191 1 : 8. 
Ghirardelli, 1950 : 109. 
Fraser, 1952 : 9. 
David, 1955 : 256. 
Furnestin, 1957 : 231. 

Sagitta lyra, S. maxima and S. gazellae have, until recently, often been confused, 
in spite of the fact that Ritter-Zahony showed that the adults can be separated on 
tail-length alone : 

Tail 
Species (as % of length) 

S. gazellae . . 10-14 

S. lyra . . . 14-18 

S. maxima . . 19-25 

There can be a certain amount of overlap between young S. lyra and old 5. maxima 
and between young 5. gazellae and old 5. lyra, since in young stages the tail tends 
to be proportionately longer ; almost all the adults, however, can apparently be 
sorted with confidence from Ritter-Zahony's figures. 

Much of the confusion has been caused by the habit of these species of shedding 
some of their hooks and posterior teeth on reaching maturity ; this habit is least 
marked in 5. maxima, most marked as regards hooks in S. lyra, and as regards 
posterior teeth in 5. gazellae. Formulae for head armatures are therefore of no value 
in these species unless the state of maturity is also given. 

David (1955) has shown that 5. gazellae is distinct from 5. lyra, not only anatomic- 
ally, but also geographically. S. gazellae is confined to the Antarctic and to the sub- 
antarctic south of the subtropical convergence, whereas S. lyra occupies subtropical 
and tropical latitudes ; the two species overlap very little, if at all. 

5. maxima is the only one of these three species in arctic waters, where it is abun- 
dant, and it also extends into the tropics and (David, 1958) to the Antarctic. It 
occurred regularly with 5. lyra in the Rosaura catches throughout the Caribbean 
and the central Atlantic. 

Furnestin (1957) has summarized observations on S. lyra by herself, Ghirardelli 
(1950) and Hamon (1952) in the general region of the Mediterranean and the water 



236 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

off north-west Africa ; these waters can be classified as warm-temperate to subtropical. 
She finds an unbroken sequence from younger forms with 6-10 hooks and 4-12 
posterior teeth, through numerous intermediates with 5-9 hooks and 8-16 posterior 
teeth, to adults with as few as 3 hooks and 2 posterior teeth ; only in the last stage 
are the ovaries well developed (= my stages III and IV). Mr. David (in a letter) 
says that in southern subtropical waters S. lyra behaves in much the same way, 
losing hooks and posterior teeth gradually with advancing maturity ; in particular 
he mentions having seen 5. lyra with 4 and 5 hooks. 

This is in contrast to the Rosaura material, in which there are either 3 hooks (one 
has only 2), or 6 or more ; in no specimens there is not one with either 4 or 5 hooks 
on either side. The difference between the two lots was so striking that at first (before 
seeing either Furnestin (1957) or David (1955)) I thought that there were two species, 
each of which identified itself as 5. lyra. Eventually, however, it became apparent 
that almost all of form A (6 hooks or more) were in stage I or II, and almost all 
form B (3 hooks) were in stages III or IV, as shown in the tables below. (I have 
divided other species into stages O and I as against stages II-IV or V, but the facts 
about 5. lyra are more clearly shown by dividing them into forms A and B as defined 
above.) 

Two of the Rosaura specimens came from Station 13 off the eastern United States, 
but all the rest were from ten tropical stations. 

Size-range and frequency of maturity stages. There were none of stage O, and I have 
included two intermediate stages, I— II and II— III. In I— II the testis was still visible 
as such but there were already free sperms in the tail ; there were two examples, 
one with 8 hooks and 10 posterior teeth, the other with 3 hooks and 3 posterior 
teeth. In stages II— III some of the eggs had increased in size as in an ovary of early 
stage III, but the sperms had not been evacuated from the tail. There were seven in 
stages II— III ; six had 3 hooks, but one had 3 hooks on one side and 6 on the other ; 
four had from 5 to 9 back teeth, one had 3, one 2, and one had 3 teeth and 6 empty 
sockets whose teeth must have been very recently lost. (Incidentally, it is the 
posterior hooks and the lateral teeth which are lost.) 

Stage 

I 

I-II 

II 

II-III 

III 

IV 

Anatomical details are given below (columns 
3 or 2 hooks). 

All but two in group B were riper than stage II, whereas none in group A was riper 
than stage II ; this shows that in the tropical Caribbean and central Atlantic the 
shedding of both hooks and posterior teeth takes place (in the great majority of 
cases) during the evacuation of sperm from the tail, i.e., while the individual is 
advancing from stage II to stage III. Furthermore, five individuals with only 3 



Size-range 
(mm.) 








Number 




13-6-26-0 








6 




i3-5,i5-8 
15-3-26-4 

I7-9-23-3 
15-0-27-5 
16-9-26-7 








2 
3i 

7 
56 

7 




(columns 


A, 


6 


or 


more hooks ; 


columns B, 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



237 











Tail 










Ai 


it. 


Post. 








Length 




(%le 


Hooks 


teeth 


teeth 


Maturity 






K 






A. 




















a 




r 




N 


r 




a 


( 




■\ 


mm. 


A 


B 


% 


A 


B 


No. 


A 


B 


A 


B 


A 


B 


Stage 


A 


B 


>27 


— 


1 


13 


1 


3 


j 


— 


— 


— 


— 


— 


1 


J 


6 


— 


25-27 


3 


11 


14 


4 


7 


2 


— 


1 


— 


— 


— 


35 


. 7-77 


1 


1 


23-25 


6 


22 


15 


10 


28 


3 


— 


7<>i 


— 


— 


— 


3i 


77 


30 


1 


21-23 


8 


18 


16 


10 


23 


4 


— 


— 


1 


— 


— 


— 


. 77-777 


— 


7 


19-21 


8 


7 


17 


7 


8 


5 


— 


— 


2 


1 


— 


1 


777 


— 


56 


17-19 


6 


5 


18 


1 


2 


6 


6 


I 


3 


11 


1 


1 


IV 


— 


7 


15-17 


5 


7 


19 


1 


1 


7 


20 


— 


13 


40 


3 


— 








<i5 


1 


1 


20 


3 




8 

9 
10 
11 


8 
3 


— 


16 

2 


20 


7 
10 

14 

2 


2 

1 









hooks still had five or more posterior teeth, whereas none in group A had fewer 
than 6 ; this shows that the hooks tend to be shed slightly in advance of the posterior 
teeth. The change-over is very abrupt in these tropical specimens, and not gradual 
(see above) as in subtropical waters. 

Apart from the length of the tail, the claw-like hooks of fully mature S. lyra 
(Ritter-Zahony, 1911, fig. 8 ; Ghirardelli, 1950, fig. 2b ; David, 1955, fig. 5c ; Fur- 
nestin, 1957, phot. 45 and 47, fig. 93) serve at once to distinguish this species from 
S. maxima. David (1955) and Fraser (1957) describe a further difference ; in 5. 
maxima the lateral nerve runs below the posterior fin, whereas in S. lyra the nerve 
splits and passes on each side of the fin. 



Sagitta maxima (Conant) 

Ritter-Z&hony, 1910 : 264 ; 191 1 : 8. 
Fraser, 1952 : 8. 

S. maxima greatly resembles S. lyra in size, transparency and flaccidity, but the 
former's tail is relatively longer and, as mentioned above, 5. maxima never develops 
the claw-shaped hooks so characteristic of mature S. lyra. 

Of the 167 specimens, 85 came from Station 8 off Greenland ; the rest were distri- 
buted fairly evenly through the remaining samples. 

5. maxima is one of the few species in the Rosaura collection to show signs of 
geographic variation, as can be seen from the tables below. As regards tail-length 
and number of anterior teeth there is agreement throughout the range covered, 
but in length, number of hooks and number of posterior teeth, the Greenland specimens 
are distinct from the tropical ones ; those from Station 13 off the eastern United 
States are intermediate, but incline more towards the Greenland forms. (Station 
13 produced 7 stage O and 5 stage I, but unfortunately no adults.) 

Length. The 42 adults (stages II-IV) from Station 8 ranged in size from 42 to 59 
mm., whereas those from Stations 27-46 ranged from 16-2 to 34-6 mm., leaving a 
gap between the two series of 7-4 mm. Admittedly, those from Station 8 were well 
preserved, whereas the others were all more or less contracted, but even so I do not 
think that the latter could ever have been nearly so large as the former. 



238 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



Among the younger specimens, all-stage I from Station 8 were larger than those 
from any other stations, most of them being as big as the adults. The O's and Fs 
from Station 13, while smaller than those from Station 8, are larger than those from 
the tropics and provide a link between the two. 

Hooks. The adults of Station 8 all possessed 6, 7 or 8 hooks, whereas those from 
the tropics had 3, 4 or 5, with only one 6 and one 7. 

The younger specimens present a less clear picture, because of their greater range 
in hook-number and because their numbers are few. 

Posterior teeth. The adults from Station 8 have more posterior teeth than do those 
from the tropics, the majority having 5-8 as compared with 3-5. Again the greater 
range in the young stages obscures the picture. 

In the following tables, adults (II-IV) are shown in heavy type. Station 8, Station 
13 and Stations 27-46 are shown separately. 



56-60 
52-56 
48-52 
44.-48 
40-44 
36-40 
32-36 
28-32 
24-28 
20-24 
16-20 
12-16 



Length 



St. 8 



0,1 II-IV 
— 5 



5 — 



St. 13 
O, I II-IV 



3 — 
6 — 
3 — 



Sagitta maxima 



Sts. 27-46 
o, I II-IV 



Tail (as % of length) 



27-46 







— — 6 

— 2 7 

— — 7 

— — 8 

— — 5 



No. 



Hooks 



St. 8 



St. 13 



Sts. 27-46 



Anterior teeth 



St. 8 



St. 13 



Sts. 27-46 



Posterior teeth 



St. 8 



St. 13 



Sts. 27-46 



O, I II-IV O, I II-IV O, I II-IV O, I II-IV O, I II-IV O, I II-IV O, I II-IV O, I II-IV O, I II-IV) 















5 


5 








6 


2 










4 












8 


16 


12 


1 


— 


3 


41 


3 


— 


— 


— 


— 


21 


— 


— 


— 


— 


1 


33 


14 


22 


9 


— 


4 


11 


7 


2 


2 


— 


2 


23 


2 


— 


— 


— 


1 


15 


5 


3 


2 


— 


1 


3 


7 


9 


4 


— 


3 


10 


4 


8 


— 


— 


4 


1 


2 


— 


— 


— 


4 


— 


15 


18 


5 


— 


2 


— 


17 

7 


24 
10 


i 
9 





I 
1 


1 


1 

















7 
2 


7 
5 


1 





1 
1 





5 
8 


— 


2 


— 


2 
2 








— 


— 


— 


— 


— 


1 


— 




2 

1 





Fowler, 1906 : n. 
Ritter-Z&hony, 191 1 : 7. 
Michael, 191 1 : 30 ; 1919 : 245 
Burfield & Harvey, 1926 : 95. 
Burfield, 1930 : 210. 
Thomson, 1947 : 10. 
Fraser, 1952 : 9. 
Vanucci & Hosoe, 1952 : 15. 
Furnestin, 1957 : 201. 



Sagitta hexaptera d'Orbigny 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



239 



This large, transparent species was not found in the samples from Stations 8 
and 13, but occurred regularly throughout the Caribbean and the central Atlantic. 
The specimens were quite well preserved (except for the fins) and were not contracted. 
The bodily proportions and head armature were very consistent throughout, and 
showed no sign of division into geographical races. The anterior teeth, which are long, 
thin, and project forwards in a very characteristic manner, afford the best diagnostic 
feature (see Furnestin (1957), fig. 81). 

Of the 299 specimens, the smallest measured 6-6 mm. (stage O) ; the largest, 
though only in stage I, measured 387 mm. 

The size-range and frequency of occurrence of the maturity stages were as follows : 















Sagitta hexapti 


zra 






















Size-range 


















Stag 


» 




(mm.) 






Number 












O 






6-6-28-2 




. 


35 












I 






11-4-38-7 




. 


133 












II 






14-0-36-6 




. 


112 












III 






15-7-28-5 




. 


15 












IV 






18-2-34-1 




• 


4 






Other anatomical details 


were 


as follows : 














Length 




Tail (% length) Hooks 




Anterior teeth 


Posterior teeth 


























mm. 


0, I II-IV Total 


% 


0,1 


II-IV Total No. 0, I II-IV Total 


O, I II-IV Total 


0,1 


II-IV Total 


36-40 


1 1 


2 : 


15 


1 


— 


1.0 — 










5 5 


32-36 


1 7 


8 


16 


— 


2 


2 


1 — 








5 


32 37 


28-32 


13 4 


17 


17 


5 


3 


8 


2 — 


— 


— 


13 16 29 


49 


60 109 


24-28 


19 28 


47 


18 


7 


7 


14 


3 ~ 


— 


— 


107 87 194 


4i 


20 61 


20-24 


38 57 


95 


19 


9 


16 


25 


4 — 


— 


— 


44 28 72 


27 


10 37 


16-20 


48 32 


80 


20 


3i 


46 


77 


5 1 


4 


5 


3 - 3 


36 


3 39 


12-16 


36 2 


38 


21 


33 


25 


58 


6 2 


10 


12 




8 


1 9 


8-12 


9 — 


9 


22 


37 


18 


55 


7 67 


44 


111 








<8 


3 — 


3 


23 


21 


8 


29 


8 32 


53 


85 














24 


12 


4 


16 


9 4i 


17 


58 














25 


10 


1 


11 


10 23 


3 


26 














26 


1 


— 


1 


11 2 




2 














27 


1 


— 


1 















The Rosaura S. hexaptera were of additional interest in that some of them contained 
two-tailed cercarias which varied in body-length from 0-35 mm. to 1-2 mm. Out of 
299 S. hexaptera 13 contained cercarias ; one was in the gut, but all the others 
were lying free in the body cavity. They came from Stations 15, 27, 32 and 33, all 
in the Caribbean Sea. Dr. Dawes has been kind enough to examine them, and 
has identified them (Dawes, 1958) as Cercaria owreae Hutton (1954). No similar 
parasites were seen in any other species, though some (e.g. 5. enflata and S. 
maxima) were quite as transparent as S. hexaptera, in which the cercarias were 
conspicuous. 

In the sample from Station 45, one S. hexaptera contained many small multi- 
nucleate parasites in the body cavity, which Dr. Dawes thinks are not helminths. 

Sagitta decipiens Fowler 
Ritter-Z&hony, 191 1 : 27. 
Michael, 1919 : 254. 
Thomson, 1947 : 20. 

The 29 specimens of this somewhat slender species came from seven stations in 
the Caribbean and central Atlantic. Twenty-one were from two stations, 27 and 28, 



240 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



in the Gulf of Honduras ; this may indicate that this species is commoner at the 
western extremities of the Caribbean Sea than it is elsewhere. 

The specimens ranged in size from 6*3 to 13-5 mm. (both were at stage II). The 
size-range and frequency of occurrence of the maturity stages were as follows : 





Stage 




Size-range 
(mm.) 


Number 








O 




7-6 




I 








I 




8-8-n-i 




6 








II 
III 




6-3-i3'5 




22 









IV 




io-8 




1 






►ther specifications were as 


follows : 










Length 
(mm.) 

A. 


Tail 
(% length) 

A_ 


Hooks 

A 


Ant. 


teeth 


Post. 


teeth 

A 


r \ 
mm. Total 


%• 


Total 


f \ 

No. Total 


No. 


Total 


No. 


Total 


12-14 8 


20 


2 


6 18 


4 


1 


10 


1 


10-12 17 


21 


10 


7 11 


5 





— 


— 


8-10 2 


22 


7 




6 


13 


13 


3 


6-8 2 


23 


5 




7 


9 


14 


4 




2 4-2g 


5 




8 


6 


15 
16 

17 
20 


6 
12 

1 
2 



Sagitta enflata Grassi David, 1956 
Fowler, 1906 : 8. 
Ritter-Z&hony, 191 1 : 13. 
Michael, 191 1 : 28 ; 1919 : 242. 
Burfield & Harvey, 1926 : 95. 
Thomson, 1947 : 11. 
Vannucci & Hosoe, 1952 : 10. 
Furnestin, 1957 : 213. 

This medium-sized, transparent species occurred throughout the Caribbean and 
the central Atlantic, and also in Station 13 off the eastern United States (it was not 
found in Station 8 off Greenland). A few very young ones (the largest measuring 
only 3-1 mm.) were caught by the small silk townet in the harbour at Belize, British 
Honduras (Station 23). This agrees with previous records that S. enflata is one of 
the few oceanic species that penetrate inshore waters ; on the Queensland coast, 
for example, it is the commonest species in the neritic waters between the Great 
Barrier Reef and the mainland (Burfield, 1950). 

In one character, the number of anterior teeth, there is a difference between those 
from the temperate Station 13 and those from the tropical stations. Of the 14 
specimens from Station 13, six had 5 anterior teeth, seven had 6 and one had 8 ; 
in the Caribbean and central Atlantic the great majority had 8 or 9, with a consider- 
able number of y's and io's. 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



241 



Of the 135 5. enflata, the smallest (excluding the very young ones mentioned above 
from Station 23) measured 5-2 mm. (stage O), and the largest was 17-1 mm. (stage II). 
The size-range and frequency of the developmental stages were as follows : 



Stage 


Size-range 
(mm.) 


Number 


O 


5-2-10-8 


9 


I 

II 

III 


7.3-15-0 

7.3-17-1 

n-5-16-5 


48 
56 
20 


IV 


8-i, 15-0 


2 



The great size-range of each stage can, in the case of 5. enflata, be explained by 
the fact that individuals of this species may breed several times, and continue to 
grow all the time (see Furnestin, 1957, p. 226). 

Other specifications, which agree with previous accounts, are as follows : 

















Sagitta enflata 


















Length 






Tail (% length) 




Hooks 




Anterior teeth 


Posteri 




mm. 


0,1 


II-IV Total 


/o 


0,1 


II-IV Total 


No. 


0,1 


II-IV 


Total 


0,1 


II-IV 


Total 


0,1 


II-IV Total 


16-18 


— 


6 


6 


13 


— 


4 


4 


5 


— 


— 


— 


6 


2 


8 


— 


— 


— 


14-16 


2 


21 


23 


14 


2 


4 


6 


6 


— 


— 


— 


1 


7 


8 


1 


— 


1 


12-14 


11 


35 


46 


15 


7 


12 


19 


7 


— 


1 


1 


5 


9 


14 


1 


— 


1 


10-12 


20 


12 


32 


16 


9 


22 


31 


8 


— 


4 


4 


17 


15 


32 


— 


— 


— 


8-10 


14 


4 


18 


17 


17 


14 


31 


9 


30 


32 


62 


21 


28 


49 


3 


— 


3 


6-8 


8 


1 


9 


18 


7 


8 


15 


10 


19 


34 


53 


6 


13 


19 


6 


5 


11 


4-6 


1 


— 


1 


19 


6 


7 


13 


11 


7 


6 


13 


— 


4 


4 


3 


3 


6 










20 


4 


2 


6 


12 


— 


2 


2 


— 


1 


1 


8 


11 


19 










21 


3 


3 


6 


13 


— 


— 


— 








8 


12 


20 










22 




3 


3 


14 


— 


— 


— 








15 


18 


33 










23 


— 


— 


— 


IS 


— 


— 


— 








4 


7 


11 










24 


1 




1 


16 

17 
18 

19 


- 


- 


- 








6 

1 


9 

8 
5 

1 


15 
9 
5 
1 



Sagitta zetesios Fowler and Sagitta planctonis Steinhaus 

David, 1956. 

These two species were for many years placed together as S. planctonis, but the 
definitive paper by David (1956) shows that they are separate, and that S. zetesios 
is much the commoner of the two. The best character for separating them is the 
number of posterior teeth in almost all specimens except juveniles ; 5. planctonis 
has less than 14, S. zetesios has more than 14. Further, in a summary of previous 
records, David states that 5. planctonis is epiplanktonic in warm water, whereas 
S. zetesios is mesoplanktonic in most deep oceans except the Antarctic. The Rosaura 
specimens fully confirm David in the matter of posterior teeth and as regards abun- 
dance and horizontal distribution (S. zetesios occurred in Stations 8 and 13 and in all 
but two of the offshore tropical stations, 5. planctonis only in Stations 45 and 46 
in the eastern central Atlantic) . Because open nets were used, however, the Rosaura 
material does not provide evidence about vertical distribution. 

David (1956) describes a third species from this group (5. marri) which is confined 
to the Antarctic, and it is perhaps worth calling attention to the remarkable paral- 
lelism in geographical distribution between this group of species and the group 



242 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

5. maxima-lyra-gazellae. Each group contains one species which is almost cosmo- 
politan (S. zetesios and S. maxima), one confined to the Antarctic (5. marri and 5. 
gazellae), and one confined to warmer waters (S. planctonis and 5. lyra.) 

S. zetesios. Length (see tables below) . The 15 specimens of S. zetesios from Station 
8 were well preserved and very large, ranging from 31 to 42-5 mm. (David (1956) 
says " Length up to 40 mm."). The tropical specimens were all contracted and the 
largest (stage II) was only 24-3 mm. long ; I doubt whether any of the tropical ones 
could have been more than about 30 mm. long when alive. Part of this discrepancy 
can be explained by the greater age and maturity of the Greenland specimens, 
but even so stage I in Station 8 reached a size of 33 mm. The few specimens from 
Station 13 were intermediate as regards length. 

Tail length. Those from Station 8 show a very narrow range of variation, all but 
three having tails 22 or 23% of the body-length. The much greater variation from 
the warmer stations is probably due to independently different degrees of contraction 
of trunk and tail. 

Hooks. Station 8 shows fewer hooks (6-9) than the others (8-1 1). This may be 
an indication of racial difference, but David (in a letter) suggests that it is more 
likely due to the greater immaturity of the tropical specimens ; it is difficult, however, 
on the latter supposition to explain away the several tropical stage II with 10 hooks, 
while no stage I from Station 8 had more than 9. The absence of stage O from 
Station 8 seems to be real. Had they been present they would have been in the same 
size-group as Eukrohnia hamata, of which I picked out 139 from Station 8. 

Anterior teeth. Here I think that the difference between Station 8 and the others 
is due solely to the maturest specimens (of which there were none from the tropics) 
having shed several teeth in the manner of S. maxima and 5. lyra. 

Posterior teeth. There seems to be no significant difference between Station 8 
and the others. 

S. planctonis. Length and maturity. In length 5. planctonis was very similar to 
the tropical S. zetesios and was about as contracted, but the largest specimens were 
much more mature. There was only one stage III S. zetesios in Stations 13-46, 
a rather small specimen of 18-3 mm.; the older ones were probably living below 
the depth fished by the nets. The epiplanktonic 5. planctonis, on the other hand, 
provided six fully matured stage III, three each of stages I and II, and no stage O. 

Tail length. This covers much the same range of variation as in the tropical 
S. zetesios. 

Hooks. David (1956) says " Hooks up to 11, usually 8-1 1 ". The few Rosaura 
specimens have 6-9, mostly 7. 

Anterior teeth. There are fewer anterior teeth (5-7, mostly 5) in 5. planctonis 
than in the tropical 5. zetesios (7-13, mostly 8-11). 

Posterior teeth. With the exception of a single very small (6-2 mm.) 5. zetesios 
with only 10 teeth, there is here an absolute distinction (confirming David, 1956) 
between the two species. In the Rosaura specimens S. zetesios have 12 posterior 
teeth or more, S. planctonis have 10 or less. 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



243 



The size-range and frequency of occurrence of maturity stages of these two species 
were as follows : 



Stage 

O 

I 

II 

III 



Sagitta zetesios 

A 



Station 8 



Station 13 
a 



Stations 27-46 



Size-range 
(mm.) 

31-33 
32'5-38 
4°>42'5 



No. 



Size-range 
(mm.) 



22 



No. 

1 



24-28-5 



Size-range 
(mm.) 

8-9-17-0 

6 • 2-20 • 5 

9-2-24-3 

18-3 



No. 

5 
33 
25 

1 



5. planctonis 
a 

Stations 45, 46 
a „ 



Size-range 
(mm.) 

9-7-18-4 
17-2-17-7 
18-9-26-9 



No. 
o 
3 
3 



Other specifications were as follows (S. zetesios on the left, 5. planctonis on the 
right) : 









Sagitta zetesios 






















a 














5. planctonis 




r 
















\ 






Station 8 


Station 13 






Stations 27-46 


Stations 45, 46 


Length 




a_ 


A. 










_A 






A 


r 


■\ 


r 


■\ 


r 






■ ^ 


c 


^ 


(mm.) 


I 


II, III Total 


O II Total 


O, I 


II 


,111 Total 


I II 


, III Total 


40-44 . 


— 


2 2 




















36-40 . 


— 


5 5 




















32-36 . 


4 


1 5 




















28-32 . 


3 


— 3 


— 1 


1 




— 






— 


— 


— — 


24-28 . 


— 


— — 


— 3 


3 




— 




1 


1 


— 


1 1 


20-24 . 


— 


— — 


1 — 


1 




2 




5 


7 


— 


3 3 


16-20 . 












6 




12 


18 


1 


5 6 


12-16 . 












16 




7 


23 


1 


— 1 


8-12 . 












1.3 




1 


14 


1 


— 1 


4-81 ■ 












1 






1 












Tail (as % 


of 


length) 


















S. zetesios 




















t 




A., 










\ 


5 


. plancton 
























Station 8 


Stations ] 


[3-46 






St 


ations 45, 


46 






, A 






>. 










a-- 






r 


^ r 








""\ 


r 




*\ 


% 




I II, III 


Total O, I 


II 


, III Total 




I 


II, III 


Total 


17 




— — 


— 3 




2 




5 




— 


2 


2 


18 




— — 


— 8 




3 




11 




— 


1 


1 


19 




— — 


— 5 




4 




9 




— 


— 


— 


20 




— — 


— 9 




6 




15 




— 


4 


4 


21 




I I 


2 3 




5 




8 




I 


1 


2 


22 




3 4 


7 3 




5 




8 




I 


1 


2 


23 




3 2 


5 2 




1 




3 




— 


— 


— 


24 




— 1 


1 3 




1 




4 




I 


— 


1 


25 




— — 


— 3 




2 




5 










26 




— — 


— — 




1 




1 











244 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

Hooks 









5. 


zetesios 
a_ 






5. planctonis 
Stations 45, 46 




t 


Station 8 




Stations 13- 


-46 






A 






-V. 




_>. 


No. 


t 
I 


II, III 


Total 


O, I 


II, III 


Total 


I II, III Total 


6 


— 


I 


1 


— 


— 


— 


— 1 1 


7 


— 


3 


3 


— 


— 


— 


7 7 


8 


3 


3 


6 


— 


4 


4 


— 1 1 


9 


4 


1 


5 


15 


17 


32 


3 — 3 


10 


— 


— 


— 


22 


9 


31 




11 


— 


— 


— 


2 


— 


2 












Anterior teeth 












S. 


zetesios 

_A_. ._ 






5. planctonis 
Stations 45, 46 




r 


Station 8 
II, III 




Qfo+irvno T ->_ 


-46 
Total 


No. 


I 


Total 


O, I 


II, III 


I II, III Total 


4 


— 


1 


1 










5 


— 


1 


1 





— 


— 


• - 7 7 


6 


— 


1 


1 





— 


— 


2 1 3 


7 


1 


1 


2 


2 


— 


2 


1 I 2 


8 


— 


3 


3 


II 


5 


16 




9 


i 


— 


1 


6 


5 


11 




10 


3 


— 


3 


15 


8 


23 




11 


2 


1 


3 


2 


8 


10 




12 


— 


— 


— 


2 


1 


3 




*3 


— 


— 


— 


1 


3 


4 












Posterior teeth 












5. 


zetesios 






S. planctonis 
Stations 45, 46 

A 




f 


Station 8 
II, III 




Qf o4*inno t o_ 


-46 
Total 


No. 


I 


Total 


OLc 

O, I 


II, III 


I II, III Total 


5 
6 














— 1 1 

— 3 3 


7 
8 














145 


9 














1 1 2 


10 


— 


— 


— 


1 





1 


I — 1 


11 
















12 


— 


1 


1 


i 





1 




13 


— 


3 


3 


2 





2 




14 


— 


1 


1 


3 





3 




15 


— 


— 


— 


2 


I 


3 




16 


4 


— 


4 


7 


3 


10 




17 
18 


3 


1 

2 


1 
5 


3 
4 


3 
5 


6 
9 




19 


— 


— 


— 


4 


9 


13 




20 


— 


— 


— 


9 


5 


14 




21 


— 


— 


— 


2 


2 


4 




22 


— 


— 


— 


1 


2 


3 





THE ROSAURA EXPEDITION T937-38: CHAETOGNATHA 



245 



Sagitta neglecta Aida 
Fowler, 1906 : 15. 
Ritter-Zahony, 191 1 : 23. 
Michael, 191 1 : 46 ; 1919 : 258. 
Thomson, 1947 : 17. 

There were seven specimens from five tropical stations (15, 23, 27 and 28 in the 
Caribbean, 46 in the eastern Central Atlantic). 



Length 


TaH 




Ant. 


Post. 




(mm.) 


(% length) 


Hooks 


teeth 


teeth 


Stage 


8-8 


25 


10 


6 


12 


III 


8-3 


24 


7 


8 


18 


II 


7-2 


25 


7 


8 


13 


II 


7-2 


26 


8 


5 


14 


I 


7.0 


27 


7 


6 


11 


I 


6-5 


22 


6 


8 


13 


I 


6-2 


3i 


7 


3 


9 


O 



Fowler, 1906 : 17. 
Ritter-Zahony, 191 1 : 21. 
Michael, 1919 : 251. 
Thomson, 1947 : 19. 



Sagitta pulchra Doncaster 



There were only two specimens, both from Station 42 off north-east Brazil. 



Length 
(mm.) 

9-6 

8-5 



Tail 
(% length) 

22 
24 



Hooks 

5 
6 



Ant. 
teeth 

8 
7 



Post, 
teeth 

16 
16 



Stage 

III 
II 



Sagitta macrocephala Fowler 
Fowler, 1905 : 65. 
Ritter-Zahony, 191 1 : 30. 
Fraser, 1952 : 10. 

This species is very easy to recognize, even with the naked eye, by its red colour 
and its large head. It is a deep-water species, and provided 107 specimens. It 
occurred off Greenland (Station 8), off the eastern United States (Station 13), in 
the Tongue of the Ocean in the Bahamas (Station 14), in two of the deepest hauls 
in the Caribbean (Stations 28 and 33), and in two of the deepest hauls in the central 
Atlantic (Stations 42 and 46). It was not picked out from the deep haul between 
Cuba and Jamaica (Station 15) ; two other regularly occurring species, S. maxima 
and 5. zetesios, were also missing from Station 15. 

The 19 specimens from Station 8 were well preserved and ranged in size from 12-5 
to 21 mm. Those from other stations were badly contracted, and did not exceed 
12-8 mm. The size-range (for what it is worth) and the frequency of maturity stages 
were as follows : 



246 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



Stage 

O 
I 

II 
III 



Size-range 
(mm.) 

5-2-11-6 
5-15-19-0 

8-I-2I'0 

20-5 



Number 

20 
60 

25 

1 



Other specifications are shown below ; S. macrocephala is remarkable for the even 
spread of the numbers of both sets of teeth in the adults, and for the range of numbers. 



Sagitta macrocephala 





Length 






Tail (% length) 






Hooks 




Anterior teeth 




Posterior teeth 


























































\ 










r ' 
















mm. 


0,1 


II, III Total 


/o 


0,1 


II, III Total 


No. 


0,1 


II, III Total 


0,1 


II 


, III Total 


\ No. 


O.I 


II, III Tot 


20-22 


— 


3 


3 


26 


1 


— 


1 . 


1 


— 


— 


— 


I 




— 


1 


16-18 


6 


— 


6 


18-20 


1 


3 


4 


27 


2 


— 


2 . 


















19 


3 


— 


3 


16-18 


— 


— 





28 


4 


1 


5 . 


4 


— 


— 


— 


— 




1 


1 


20 


5 


— 


5 


14-16 


7 


1 


8 


2 9 


5 


1 


6 . 


5 


— 


— 


— 


3 




2 


5 


21 


9 


— 


9 


12-14 


7 


2 


9 


30 


5 


3 


8 . 


6 


— 


— 


— 


14 




3 


17 


22 


4 


— 


4 


10-12 


5 


7 


12 


31 


7 


2 


9 . 


7 


— 


— 


— 


35 




8 


43 


23 


3 


1 


4 


8-10 


18 


10 


28 


32 


16 


1 


17 . 


8 


— 


— 


— 


14 




4 


18 


24 


8 


3 


11 


6-8 


34 


— 


34 


33 


11 


6 


17 . 


9 


— 


— 


— 


9 




5 


14 


25 


4 


2 


6 


4-6 


8 


— 


8 


34 


7 


3 


10 . 


10 


— 


1 


1 


3 




3 


6 


26 


8 


— 


8 










35 


8 


2 


10 . 


11 


16 


4 


20 










27 


7 


2 


9 










36 


5 


1 


6 . 


12 


44 


14 


58 










28 


5 


4 


9 










37 


5 


1 


6 . 


13 


20 


7 


27 










29 


8 


4 


12 










38 


3 


2 


5 


















30 


3 


1 


4 










39 


1 


1 


2 


















3i 


2 


1 


3 










40 




2 


2 


















32 
33 
34 
35 
37 


1 
1 
2 


2 
2 
2 

1 


3 
3 

4 
1 

1 



Pterosagitta draco (Krohn) 

Ritter-Zahony, 191 1 : 33. 
Michael, 1 919 : 264. 
Ghirardelli, 1950 : 121. 
Furnestin, 1957 : 2 4 6 - 

This small, easily recognized species was taken in small numbers at almost every 
station in the Caribbean and central Atlantic. According to Burfield & Harvey 
(1926) and Thomson (1947) it lives mainly near the surface ; accordingly it would 
have beeen encountered by the Rosaura's nets only during the last few minutes of 
each haul, and in any case it is too small to be caught reliably by a stramin net. 
These considerations indicate, I think, that P. draco must be common or abundant 
throughout the tropical waters sampled by the Rosaura. 

Burfield & Harvey (1926) record P. draco as abundant in the Indian Ocean, 
Burfield (1930) as abundant in the central Atlantic, Tokioka (1939, 19400) as common 
in the warmer waters off Japan, Thomson (1947) as common to the south-east 
of Australia, and Pierce (1953) as common off North Carolina. Nevertheless Furnestin 
(1957) states : " les auteurs s'accordent a signaler sa rarete, non seulement en 
Mediterranee mais aussi dans TAtlantique et le Pacifique (exception faite pour 
Thomson, qui la classe parmi les especes ' sub-dominantes ' au sud-est de TAustralie." 

There are 45 specimens in the Rosaura collection, ranging in size from 4-4 mm. 
(stage II) to 7-8 mm. (stage III) ; many of them are rather contracted. The size- 
range and frequency of occurrence of the maturity stages are as follows : 



THE ROSAURA EXPEDITION 1937-38 


: CHAETOG 


NATHA 










Size-range 










Stage 




(mm.) 


Number 








O 




4-5.4-7 


2 








I 




4 • 85-6 • 25 


8 








II 




4-4-7*45 


14 








III 




5-5-7'8 


18 








IV 




5-4, 6-5. 7*° 


3 






Other specifications are as 


follows 












Tail 










Length 


(% length) 


Hooks 


Ant. teeth 


Post. 


teeth 


A 


a 




A 


a 




A 


r \ 


r 


*\ 


r \ 


r ^ 


r 


■\ 


mm. Total 


% 


Total 


No. Total 


No. Total 


No. 


Total 


7-8 11 


31-38 


5 


8 2 . 


6 11 


10-13 


3 


6-7 17 . 


39 


4 


9 10 


7 13 . 


14 


9 


5-6 13 . 


40 


15 


10 28 


8 16 


15 


10 


4-5 4 . 


4i 


5 


JJ 4 . 


9 4 . 


16 


8 




42 


5 


12 1 




17 


6 




43 


5 






18 


7 




44 


3 






19 


1 




45-47 


3 






22 


1 



247 



Krohnitta subtilis (Grassi) 
Michael, 191 1 : 52. 
Tokioka, 1939 : 135. 
Thomson, 1947 : 22. 
Vannucci & Hosoe, 1952 : 25. 

The 11 specimens of this very slender species were all from tropical stations. Several 
authors have attempted to join K. pacifica with this species, but the descriptions of 
Tokioka (1939) and Thomson (1947), make it certain that the two species are distinct. 
Dr. Pierce kindly sent me a sample of his Floridan K. pacifica, which was a great 
help in confirming that all the Rosaura specimens do belong to K. subtilis. This is 
of some interest, in that both Tokioka in Japanese waters, and Pierce (1951, 1953) 
off Florida and North Carolina, stress the comparative scarcity of K. subtilis as 
compared with K. pacifica. 

The details of the Rosaura specimens were as follows (there is only one row of 
teeth in Krohnitta) : 



.ength 


Tail 








mm.) 


(% length) 


Hooks 


Teeth 


Stage 


n-8 


36 


8 


11 


II 


n-3 


33 


8 


11 


II 


10-7 


34 


9 


12 


III 


10-7 


38 


9 


10 


II 


io-6 


32 


9 


12 


III 


io-6 


38 


9 


10 


III 


100 


35 


8 


10 


III 


io-o 


33 


9 


11 


III 


9-8 


33 


8 


12 


II 


9*25 


39 


8 


10 


II 


90 


36 


9 


9 


O 



248 



THE ROSAUBA EXPEDITION 1937-38: CHAETOGNATHA 



Eukrohnia hamata (Mobius) 

Ritter-Zahony, 1910a : 268 ; 191 1 : 39. 
Michael, 191 1 : 39. 
Fraser, 1952 : 10. 

As recently as 1947 Thomson stated that E.fowleri was synonymous with E. hamata, 
but there can be no doubt that, as most authors have maintained, the two species 
are separate. In E. hamata the eyes usually lack black pigment, the gut is not as 
a rule coloured red, and few, if any, have more than 11 hooks. In E. fowleri the eyes 
have a conspicuous patch of black pigment, the gut is a bright red, and few, if any, 
have fewer than 12 hooks. It seems probable that Thomson did not have any 
E. fowleri (a deep-living species) in his material. 

As has been mentioned earlier, the Rosaura chaetognaths were sorted immediately 
after capture into red and colourless forms, and this served to separate 275 red 
E. fowleri from 204 E. hamata with only a single doubtful case. All the E. hamata 
had 7, 8, 9 or 10 hooks, whereas all but one of the E. fowleri had 12, 13 or 14 ; the 
exception was a very small (7-5 mm., stage O) E. fowleri with only 10 hooks, but this 
individual was too young to have developed its full complement of hooks and so does 
not affect the argument. Not a single one of either species had 11 hooks on either side. 

There were 204 specimens of E. hamata, of which 139 were in the sample off 
Greenland (Station 8) agreeing with Kramp (1939) who found E. hamata abundant 
in these waters. It was again the most abundant chaetognath from Station 13 off 
the eastern United States, but after that there were only one between Cuba and 
Jamaica (Station 15), seven off NE. Brazil (Station 42) and six south-west of Dakar 
(Station 46), these last three being relatively deep tropical hauls. I found none in 
the Caribbean Sea. E. hamata has been recorded at the surface in both northern 
and southern high latitudes, but in low latitudes only at depths where the temperature 
stays below about 12 C. (Fowler, 1906 ; Ritter-Zahony, 191 1 ; Johnston & Taylor, 
1921 ; Burfield & Harvey, 1926 ; Burfield, 1930 ; Kramp, 1939 ; Thomson, 1947). 

In Eukrohnia, in contrast to Sagitta, " spents " are not uncommon in which the 
ovaries remain visible as shrunken remnants ; following Kramp (1939) I have 
classified these " spents " as stage V. 

As in the case of Sagitta maxima (q.v.) there seems to be some correlation between 
size and latitude. In Station 8 (off Greenland) the largest specimen measured 34 mm., 
in Station 13 (off the eastern United States) 25-2 mm., and in the tropical stations 
16-1 mm. The tropical specimens were somewhat more contracted than the others, 
but never so much so as to make it possible that they were as large when alive. 

The size-range and frequency of occurrence of the maturity stages were as follows : 





Station 8 




Station 13 




Stations 15, 


42, 46 




(Arctic) 

A 




(Temperate 
a 


) 


(Tropical) 

A 




f 


^\ 


( 


^ 


r 


> 




Size-range 




Size-range 




Size-range 




Stage 


(mm.) 


No. 


(mm.) 


No. 


(mm.) 


No. 


O 


9-5-i9'5 


21 


11-2-23-0 


12 


— 


— 


I 


16-5-27-5 


25 


10-7-24-5 


25 


7-2 


1 


II 


20-0-31-0 


52 


18-7-24-2 


9 


94, 16 • 1 


2 


III 


17-0-29-0 


18 


21-8-25-1 


4 


8-8-H-2 


3 


IV 


20-5-34*0 


18 


— 


— 


9-5 


1 


V 


20-5-29-0 


5 


25-2 


I 


13-3 


1 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



249 



Other specifications are shown below. (As with Krohnitta, Eukrohnia possesses 
only one row of teeth, but whereas those of Krohnitta appear to be homologous with 
the anterior teeth of Sagitta, those of Eukrohnia are clearly homologous with the 
posterior teeth of Sagitta.) 

Eukrohnia hamata 





Length 




Tail (% 


of length) 




Hooks 






Teeth 
























A 








A 




l 






•\ 


t 






•\ 


( 






•\ 


r 








mm. 


O, I 


II-V 


Total 


% 


O, I 


II-V 


Total 


No. 


O, I 


II-V 


Total 


No. 


0,1 


II-V 


Total 


32-36 


— 


1 


1 


20 


1 


— 


1 


7 


— 


I 


1 


3-9 


14 


I 


15 


28-32 


— 


16 


16 


21 


4 


3 


7 . 


8 


19 


17 


36 


10 


3 


I 


4 


24-28 


23 


49 


72 


22 


7 


7 


14 . 


9 


58 


87 


146 


11 


4 


I 


5 


20—24 


13 


39 


52 


23 


7 


10 


17 . 


10 


7 


9 


16 


12 


5 


— 


5 


16-20 


21 


3 


24 


24 


7 


14 


21 










13 


2 


I 


3 


12-16 


20 


1 


21 


25 


8 


6 


14 










14 


1 


— 


1 


8-12 


6 


5 


11 


26 


11 


13 


24 










15 


4 


I 


5 


4-8 


1 


— 


1 


27 


6 


14 


20 










16 


— 


3 


3 










28 


9 


18 


27 










17 


1 


5 


6 










29 


7 


18 


25 










18 


9 


18 


27 










30 


5 


3 


8 










19 


13 


18 


31 










31 


6 


4 


10 










20 


12 


15 


27 










32 


3 


1 


4 










21 


3 


15 


18 










33 


2 


2 


4 










22 


7 


17 


24 










34 


1 


— ■ 


1 










23 


1 


12 


13 










35 


— 


1 


1 










24 
25-28 


2 
3 


3 
3 


5 
6 



Eukrohnia fowleri Ritter-Zahony 

Ritter-Zahony, 191 1 : 40. 
Fraser, 1952 : 10. 

The three main criteria for separating E. fowleri from E. hamata (colour, number of 
hooks, eye pigment) have been discussed above in the section on E. hamata. 

Both species occurred together in several of the Rosaura hauls, but their relative 
abundance varied greatly. In the two most northerly stations (8 and 13) E. fowleri 
was much less abundant than E. hamata, but in the tropical stations it was much 
more so, and was, indeed, the commonest chaetognath in several of the samples. 
It was not found in every tropical haul, being absent from those where the estimated 
depth of the net was less than 900 m., and common only when the net went deeper 
than 1,000 m. E. fowleri is widely recognized as an inhabitant of considerable 
depths ; in this respect it resembles the other red species, Sagitta macrocephala, 
whose incidence in the Rosaura collection was strikingly similar. 

As in the case of E. hamata, E. fowleri reached a larger size in higher latitudes 
than in the tropics, but the difference was not so marked. 

There were altogether 275 specimens of E. fowleri, of which all but 20 were caught 
in the tropics. The smallest was the 7-5 mm. juvenile with but 10 hooks already 
mentioned (p. 248) from the Gulf of Honduras (Station 28), and the largest measured 
29-5 (stage I, Station 8). 



250 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 



The size-range and frequency of occurrence of the maturity stages were as follows 
Stations 8 and 13 





(Arctic and Temperate) 


Tropical stations 






Size-range 


Size-range 


Total 


Stage 


(mm.) 


(mm.) 


number 


O 


— 


7-5-10-8 


5 


I 


16-9-29-5 


8-2-21-2 


114 


II 


25-2-28-0 


I4-3-23-0 


67 


III 


24-2-28-7 


17-6-24-7 


36 


IV 


— 


I8-0-2I-6 


5 


V 


27-0 


17-6-27-8 


38 



Other specifications were as follows : 



Eukrohnia fowleri 





Length 






Tail (% length) 






Hooks 






Teeth 






A 
















A 








-A. 




( 




•\ 


( 






% 


1 






■\ 


r 






- 1 


mm. 


O, I II-V 


Total 


/o 


O, I 


II-V 


Total 


No. 


O, I 


II-V 


Total 


No. 


O.I 


II-V 


Total 


28-32 


2 3 


5 


17 


2 


5 


7 


10 


1 


— 


1 


10—15 


24 


— 


24 


24-28 


2 16 


18 


18 


9 


6 


15 


11 


— 


— 


— 


16 


3 


— 


3 


20-24 


6 60 


66 


19 


12 


12 


24 


12 


52 


49 


102 


17 


10 


3 


13 


16—20 


36 62 


98 


20 


22 


14 


36 


13 


64 


89 


153 


18 


7 


2 


9 


12-16 


5i 5 


56 


21 


3i 


26 


57 


14 


1 


9 


10 


19 


12 


4 


16 


8-12 


19 — 


19 


22 


16 


3i 


47 










20 


8 




9 


4-8 


3 — 


3 


23 


9 


21 


30 










21 


6 


2 


8 








24 


10 


9 


19 










22 


16 


11 


27 








25 


4 


12 


16 










23 


10 


16 


26 








26 


2 


4 


6 










24 


10 


18 


28 








27 


1 


2 


3 










25 


6 


23 


29 








28 


1 


3 


4 










26 


4 


24 


28 








29 




1 


1 










27 
28 

29 
30 
31 
32 


3 


20 

11 

5 

5 


23 
11 

5 
5 
























— 


2 


2 



REFERENCES 
Aida, T. 1897. Chaetognaths from Misaki Harbour. Annot. Zool. Japon, 1. 
Bieri, R. 1957. The Chaetognath Fauna off Peru in 1941. Pacific Science, 11 : 255-264. 
Burfield, S. T. 1930. Chaetognatha. Brit. Antarctic (" Terra Nova ") Exp. 19 10. Zoology, 
7 : 203-228. 

1950. Chaetognatha. Sci. Rep. Great Barrier Reef Exp. 5, 459-473. 

& Harvey, E. J. W. 1926. The Chaetognatha of the " Sealark " Expedition. Trans. 

Linn. Soc. Lond., Ser. 2, Zoology, 19 : 93-119. 

Clarke, G. L., Pierce, E. L. & Bumpus, D. F. 1943. The distribution and reproduction of 

Sagitta elegans on Georges Bank in relation to the hydrographic conditions. Biol. Bull. 

85 : 201-226. 
Colman, J. S. 1954. The " Rosaura " Expedition. 1. Gear, Narrative and Station List. 

Bull. Brit. Mus. (nat. Hist.) Zool. 2 : 1 19-130. 
Colman, J. S. & Cooper, L. H. N. 1954. The " Rosaura " Expedition. 2. Under-water 

illumination and ecology in tropical estuaries. Ibid. 2 : 1 31-138. 
Conant, F. S. 1896. Notes on the Chaetognaths. Johns Hopkins Univ. Circ. 15 : 82-85. 
*Costa, A. 1869. Di un nuovo genere di Chetognati. Ann. Mus. zool. Univ. Napoli, 

5 : 54-57- 



THE ROSAURA EXPEDITION i937~38: CHAETOGNATHA 



251 



Summary of Tail-lengths and Head Armatures as Shown by the Great Majority of 
Adults (Stages II and up) in the Rosaura Collection 







Length 




Tail 




Hooks 




Ant. teeth 


Post, teeth 


Species 




(mm.) 


(as 


% length) 


(No.) 




(No.) 




(No.) 


Sagitta friderici 




12-7 




26 




8 




8 




18 


S. tenuis . 




4-6 




27-33 




8,9 




4-7 




10-14 


S. bipunctata 




6-12 




24-28 




8-10 




6,7 




13-16 


S. robusta 




9-12 




26 




7 




8-10 




12 


S. serratodentata 




7-10 




22-26 




6,7 




8-10 




15-22 


S. lyra : 






















Stages I, II 
Stages III, IV 


} 


15-27 




14-17 


{ 


6-8 
3 


} 


6-8 


{ 


8-10 

2,3 


S. maxima : 






















Arctic . 
Tropics 




40-60 
16-35 


} 


20-27 


{ 


6-8 
3-5 


} 


2-4 


{ 


5-8 
2-5 


S. hexaptera 




16-36 




18-23 




6-9 




2-4 




1-4 


S. decipiens 


. 


10-14 




21-23 




6,7 




6-8 




13-16 


S. enflata 


. 


7-17 




13-20 




9, 10 




6-10 




10-18 


S. planctonis 


. 


16-25 




17-22 




7 




5 




6-8 


S. zetesios : 






















Arctic . 


. 


32-44 




21-24 




7-9 




4-1 1 




12-18 


Temperate and 


Tropics 


10-28 




17-25 




8-10 




8-13 




15-22 


S. neglecta 




7-9 




24.25 




7, 10 




6,8 


• 


[2, 13, l8 


S. pulchra 




8-10 




22, 24 




5,6 




7,8 




16 


S. macrocephala 




8-22 




30-40 




11-13 




5-10 




24-34 


Pterosagitta draco 




4-8 




39-44 




9-1 1 




<H> 




14-18 


Krohnitta subtilis 


. 


9-12 




32-39 




8,9 




10-12 







Eukrohnia hamata 




12-32 




22-29 




8-10 




— 




16-24 


E. fowleri 


• 


12-32 




17-26 




12-14 




— 




22-30 



David, P. M. 1955. The distribution of Sagitta gazellae Ritter-Zahony. Discovery Rep. 

27 : 235-278. 
1956. Sagitta planctonis and related forms. Bull. Brit. Mus. (nat. Hist.) Zool. 4 : 435- 

451. 



1958. The distribution of the Chaetognatha of the Southern Ocean. Discovery Rep. 

29 : 199-228. 

Dawes, B. 1958. Sagitta as a host of larval trematodes, including a new and unique type 

of cercaria. Nature, 182, 960-1. 
Doncaster, L. 1902. Chaetognatha. Fauna & Geog. Maid. 6- Lace. Archip. 1 : 209-218. 
Faure, Marie-Louise. 1953. Contribution a l'etude morphologique et biologique de deux 

chaetognathes des eaux atlantiques du Maroc : Sagitta friderici Ritter-Zahony et Sagitta 

bipunctata Quoy et Gaimard. Vie et Milieu, 3 : 25-43. 
Fowler, G. H. 1905. Biscayan plankton collected during a cruise of H.M.S. " Research ", 

1900 ; part III : The Chaetognatha. Trans. Linn. Soc. Lond., Ser. 2, Zool. 10 : 55-87. 

1906. The Chaetognatha of the " Siboga " Expedition. Siboga Exp. 21 : 1-86. 

Fraser, J. H. 1952. The chaetognatha and other zooplankton of the Scottish area and 

their value as biological indicators of hydrographical conditions. Scot. Home Dept. Mar. 
Res. 1952, 2 : 1-52. 

J 957- Chaetognatha. Fiches Ident. Zoopl. 1 (first revision) : 1-6. 



252 THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 

Furnestin, Marie-Louise. 1954. Interet de certain details anatomiques, jusqu'ici peu 
etudies, pur la determination des chaetognathes. Bull. Soc. Zool. France, 79 : 109-112. 

1957- Chaetognathes et zooplancton du secteur atlantique marocain. Rev. Trav. Inst. 

Peches marit. 21 : 1-356. 

Ghirardelli, E. 1950. Osservazioni biologiche e sistematiche sui chetognathi della Baia 
Villefranche sur Mer. Boll. Pesca, Pisicolt. Idrobiol., Anno 26, 5 (n.s.) : 105-127. 

1951- Cicli di maturita sessuale nelle gonadi di 5. inflata Grassi del Golfo di Napoli. 

Boll. Zool. 18 : 4-6. 

1952. Osservazioni biologiche e sistematiche sui chetognathi del Golfo di Napoli. 

Pubbl. Staz. Napoli, 23 : 296-312. 

Hamon, M. 1952. Note complementaire sur les chaetognathes de la Baie d'Alger. Bull. 

Soc. Hist. nat. Afr. Nord, 43 : 50-52. 
Hosoe, K. 1956. Chaetognaths from the Isles of Fernando de Noronha. Oceanografia 

Biologica, 3 : 1-9. 
Huntsman, A. G. 1919. Biology of Atlantic waters of Canada. Some quantitative and 

qualitative studies of the eastern Canadian plankton. 3. A special study of the Canadian 

chaetognaths, their distribution etc., in the waters of the eastern coast. Canad. Fish. Exp., 

1914-15. Dept. Naval Sci. 421-485. 
Hutton, R. F. 1954. Metacercaria owreae, n. sp., an unusual trematode larva from Florida 

Current chaetognaths. Bull, mar Sci. Gulf &• Caribbean, 4 : 104-109. 
Johnston, T. H. & Taylor, B. B. 192 i. The chaetognatha. Australian Ant. Exp. 1911-14 ; 

Sci. Rep. Ser. C 6, pt. 2 : 1-16. 
Kramp, P. L. 1917. The chaetognatha collected by the " Tjalfe " Expedition to the west 

coast of Greenland in 1908 and 1909. Vid. Medd., Dansk. Naturh, Foren. 69 : 17-55. 

1939. The Godthaab Expedition, 1928 : Chaetognatha. Medd. Grenld. 80, nr. 5 : 

3-4°- 
Krohn, A. 1853. Nachtragliche Bemerkungen uber den Bau der Gattung Sagitta. Arch. 

Naturg. 19 (1) : 266-281. 
Michael, E. L. 1908. Notes on the identification of the chaetognatha. Biol. Bull. 15 : 

67-84. 

191 1. Classification and vertical distribution of the chaetognatha in the San Diego 

region. Univ. Calif. Pub. Zool. 8 (3) : 21-186. 

191 9. Report on the chaetognatha collected by the United States fisheries steamer 



" Albatross " during the Philippine expedition, 1907-1910. U.S. nat. Mus. Bull. 100 

(i(4)) •• 235-277. 
*M6bius, K. 1875. Die Expedition zur physikalisch-chemichen und biologischen Unter- 

suchung der Nordsee im Sommer, 1872. Wiss. Meerestunters. Kiel, Jahrg. 2. 
Orbigny, A. d'. 1843. Voyage dans VAmerique meridionale execute dans le cours des annees 

1826-1833. Vol. V (3) : 140-144. Paris. 
Pierce, E. L. 195 1. The chaetognatha of the west coast of Florida. Biol. Bull. 100 : 

206-228. 

1953- The chaetognatha over the continental shelf of North Carolina with attention 

to their relation to the hydrography of the area. /. mar. Res. 12 : 75—92. 

1958. The Chaetognatha of the inshore waters of North Carolina. Limnol. <S> Oceanog. 

3 : 166-170. 

*Quoy, J. & Gaimard, P. 1827. Observations zoologiques faites a bord de l'Astrolabe, en Mai 

1826, dans le detroit de Gibraltar. Ann. Sc. nat. 10 : 5-239. 
*Ritter-Zahony, R. v. 1909. Die Chatognathen der Gazelle-Expedition. Zool. Anz. 34. 

1910a. Die Chatognathen. Fauna Arctica, 5 (1) : 249-288. 

1910&. Westindische Chatognathen. Zool. Jahrb. Suppt. 11 : 133-144. 

191 1. Revision der Chatognathen. Deutsche Sudpol.-Exp. 1901-03, Bd. 13 ; Zool. 

Bd. 5 : 1-72. 

Russell, F. S. 1932a. On the biology of Sagitta. The breeding and growth of Sagitta 
elegans Verrill in the Plymouth area, 1930-31. /. mar. biol. Ass. U.K. 18 : 131-145. 



THE ROSAURA EXPEDITION 1937-38: CHAETOGNATHA 253 

Russell, F. S. 19326. On the biology of Sagitta. II. The breeding and growth of Sagitta 
setosa J. Muller in the Plymouth area, 1930-31, with a comparison with that of 5. elegans 
Verrill. Ibid. 18 : 147-160. 

1933a. On the biology of Sagitta. III. A further observation on the growth and 

breeding of Sagitta setosa in the Plymouth area. Ibid. 18 : 555-558. 

19336. On the biology of Sagitta. IV. Observations on the natural history of Sagitta 



elegans Verrill and Sagitta setosa J. Muller in the Plymouth area. Ibid. 18 : 559-574. 
Tchindinova, Y. G. 1955. Chaetognatha of the Kurile-Kamchatka trench. Trudy Inst. 

Okeanolog. 12 : 298-310 (in translation). 
Thomson, J. M. 1947. The chaetognatha of south-eastern Australia. Coun. sci. ind. Res. 

(Div. Fish. Rep. 14) Bull. 222 : 1-43. 
Tokioka, T. 1939. Chaetognaths collected chiefly from the bays of Sagami and Suruga, 

with some notes on the shape and structure of the seminal vesicle. Rec. Oceanog. Works 

Japan, 10 (2) : 123-150. 

1940a. The chaetognath fauna of the waters of western Japan. Ibid. 12 : 1-22. 

19406. A small collection of chaetognaths from the coast of New South Wales. Rec. 

Austral. Mus. 20 (6) : 367-379. 

1955. Notes on some Chaetognaths from the Gulf of Mexico. Bull. mar. Sci. Gulf 6- 



Caribbean, 5 : 52-65. 
Vannucci, K. & Hosoe, K. 1952. Resultados cientificos do cruziero do " Baependi " e do 

" Vega " a Ilha da Trinidade : Chaetognatha. Bol. Inst, oceanog. Sao Paulo, 3 : 3-34. 
Van Oye, P. 193 1. La fecondation chez les chaetognathes. Bull. Mus. roy. Hist. nat. Belg. 
7 : 1-7. 
♦Verrill, A. E. 1873. Report on the invertebrate animals of Vineyard Sound. Rep. U.S. 
Fish. Comm. 1. 

(* Not seen.) 




PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 

BARTHOLOMEW PRESS, DORKING 



A NORTH BORNEAN 

PYGMY SQUIRREL, 

GLYPHOTES SIMUS THOMAS, 

AND ITS RELATIONSHIPS 



J. E. HILL 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 9 

LONDON: 1959 



A NORTH BORNEAN PYGMY SQUIRREL, 

GLYPHOTES SIMUS THOMAS, AND ITS 

RELATIONSHIPS 



BY 

J. E. HILL 

Department of Zoology, British Museum (Natural History) 



Pp. 255-266 ; Plates 7-8 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 9 

LONDON: 1959 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
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within one calendar year. 

This paper is Vol. 5, No. 9 of the Zoological 
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Issued February, 1959 Price Seven Shillings 



A NORTH BORNEAN PYGMY SQUIRREL, 

GLYPHOTES SIMUS THOMAS, AND ITS 

RELATIONSHIPS 

By J. E. HILL 

Only three specimens of this unusual Pygmy Squirrel, Glyphotes simus, have been 
recorded since publication of the description in 1898. It has received little attention 
in the literature, no doubt through the paucity of the available material, and has 
remained little known. The British Museum (Natural History) has recently received 
a specimen of Glyphotes from the Institute for Medical Research, Kuala Lumpur, 
and this accession has prompted further examination of the genus. 

I am indebted to J. L. Harrison, lately of the Institute for Medical Research, who 
arranged the gift of Glyphotes ; to C. A. Gibson-Hill, Director, Raffles Museum, 
Singapore and to Tom Harrisson, Curator, Sarawak Museum, Kuching, who 
courteously loaned the material of Glyphotes in their care for comparison ; lastly, 
to R. W. Hayman, of the British Museum (Natural History), for his valuable advice 
and criticism during the preparation of this paper. 

Apart from the original description, incidental mention in faunal works such as 
that of Banks (193 1) and enumerations of the principal characters such as that of 
Ellerman (1940), previous published work on the genus appears limited to that of 
Chasen & Kloss (1927). Their paper gives a brief account of the known specimens 
at that date, with measurements, and records an additional specimen. 

The following account is based on five specimens of Glyphotes simus, which so far 
as I have been able to ascertain represent the total preserved in collections at the 
present time. They are : 

(i) British Museum 98. 11. 3. 6. The type specimen. Adult, sex unknown. 
Collected on Mount Kina Balu, North Borneo, by A. Everett. Skin and skull, 
the rear of the cranium missing. 

(ii) Sarawak Museum 55.11. Adult male. Collected on Mount Merapok (Mount 
Marabok), Brunei, by J. Waterstraat in December, 1899. Skin and skull, the latter 
not available : the Sarawak Museum collection was disorganized during the Japanese 
Occupation, 1941-45. Formerly British Museum 0.10.8.2 but given in exchange 
to the Sarawak Museum in 190 1. As remarked by Chasen & Kloss (1927) the label 
has been marked " Co-type of the species ", However, as these authors point out, 
the specimen was collected in the year following publication of the description and 
no mention is made by Thomas of any specimen other than the type. 

(iii) Sarawak Museum 55.12. Adult female. Collected on Mount Kina Balu, 
North Borneo at 3,500 ft. by a native collector on 10th September, 1913. Skin 

ZOOL. 5, 9. I2§ 



258 A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 

and skull : the latter not available (see (ii)). Formerly referred to Callosciurus 
notatus, which possibly explains why it was overlooked by Chasen & Kloss (1927). 

(iv) Raffles Museum. Subadult female. Collected at Tenompok, Kota Bellud, 
near the south foot of Mount Kina Balu, North Borneo, by a native collector on 10th 
June, 1925. Skin and skull, the latter much broken and fragmentary. 

(v) British Museum 58.446. Adult male. Collected at Tenompok, Mount Kina 
Balu, North Borneo, by a collector of the Institute for Medical Research, Kuala 
Lumpur, on 16th June, 1952. Specimen preserved in alcohol, the skull and baculum 
extracted and cleaned. 

GLYPHOTES Thomas 

1898 Glyphotes Thomas, Ann. Mag. nat. Hist. 2 : 250. 

A genus of dwarf squirrel characterized by its short, broad rostrum, much reduced 
postorbital processes and peculiar, specialized incisors. The zygomatic plate slants 
upwards and is more nearly vertical than in Callosciurus. The lower jaw is weak, 
with low coronoid processes and slender, elongated condylar processes. The upper 
incisors are very broad and yet not thickened antero-posteriorly, their tips divergent 
from each other. The lower incisors are similar but are more strongly divergent. 
Cheek teeth 5/4, with their pattern as in Callosciurus. 



Glyphotes simus Thomas 

1898 Glyphotes simus Thomas, Ann. Mag. nat. Hist. 2 : 250. Mount Kina Balu, North 
Borneo. 

Externally characterized by its deep, broad, short muzzle. A specimen in alcohol 
confirms the " stumpy-nosed " description applied by Thomas and indicates more 
than do study skins the depth of the muzzle in relation to its length. The ear is 
of the Callosciurine type and when laid forward reaches almost to the posterior 
can thus of the eye. The feet are of the normal tree squirrel pattern : of the four 
digits of the manus, D4 is slightly longer than D3 while the outer digits are equal in 
length and are about half the length of D4. The hallux is the shortest digit of the 
tarsus while D5 is considerably longer, nearly as long as D4, which only slightly 
exceeds in length D2 and D3. The claws are well developed, white tipped and dark 
brown at the base. The tail is rather narrow and is shorter than the head and 
body. External measurements (in millimetres) of B.M. 58.446: head and body 
115, tail 95, hindfoot 27, ear 13. 

The dorsum is dull grey, grizzled and flecked with pale buff. The dorsal pelage 
is typically of Callosciurine type, the short hairs of the underfur slate grey at the 
base and tipped with pale buff or straw colour. The longer, overlaying bristle 
hairs are black or chestnut based and are tipped with chestnut, with a subapical 
annulation of buff : not infrequently bristle hairs are found with dark tips and 
bases but with two annulations of buff separated by a central band of black. 
Occasionally, these hairs are black throughout their length. Essentially similar, 
but slightly shorter pelage extends over the nape and crown. The outer surfaces 



A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 259 

of the legs and the dorsal surfaces of the fore feet are the same colour as the back 
while the dorsal surfaces of the hind feet are slightly more buffy. The bases of the 
claws are furnished with a small tuft of brighter, more buffy hairs. The sides of 
the muzzle are brighter buff than the back and there is an orange buff eye ring. The 
ears, especially on their posterior edge, are fringed with bright orange buff. There 
is a very indistinct post auricular patch of white. The white lateral flank stripes 
are broad and are usually the same width throughout their length : in the type 
specimen they are broader anteriorly than posteriorly. The black sublateral stripes 
are narrower and less prominent, more diffuse, especially posteriorly, where a 
suffusion of black tends to spread across the belly. The anterior part of the ventrum 
is dull orange buff : posteriorly the ventrum is buffy but is suffused with grey and 
overlaid slightly with black. The tail is blackish, grizzled with bright orange buff. 
The hairs are dark tipped and dark based with a broad sub-basal annulation of 
bright buff, sometimes with a second subterminal annulation of paler buff separated 
from the sub-basal annulation by a band of black. 

The specimen from Mount Merapok, Brunei, is brighter and less greyish dorsally 
than those from Kina Balu. Its tail hairs are annulated with brighter orange and 
the underparts are more buffy and less suffused with grey, while the dorsal surfaces 
of the feet are warmer in tone. This circumstance was noted by Chasen & Kloss 
(1927), who compared this specimen with that listed as (iv) above. 

Externally, Glyphotes simus is similar to Callosciurus notatus dulitensis, from which 
it may be distinguished by smaller size, foreshortened muzzle, paler underparts and 
feet, broader, more prominent white lateral stripes, less prominent black sublateral 
stripes, greyer back and darker tail. Dorsally, it is similar in colour to Callosciurus 
nigrovittatus orestes but is paler and less buff. The annulations of the tail hairs are 
brighter than in orestes while the underparts are predominantly buff and not grey. 

The baculum of Glyphotes simus (Plates 8 (c)-8 (/)) is of the Callosciurine pattern, 
as described by Thomas (1915) and figured by Didier (1952). It consists of two 
parts, a slender, tapering shaft and a separate plate or lamina attached to its dorsal 
surface. 1 The shaft, which at its base and for most of its length is greater in depth 
than in width, is comparatively broad based and tapers only slightly in its proximal 
half, but beyond its centre point tapers more abruptly to a blunt point furnished 
laterally with two small barbs. Viewed laterally, the ventral face forms a compound 
curve, the proximal half concave, the distal half convex. The dorsal surface is 
concave, more especially in its distal portion, whereon is attached the separate 
lamina. Viewed dorsally, the baculum is a slender, straight shaft. The lamina 
originates from a point just posterior to the tip of the shaft and extends along the 
dorsal surface for about half the length of the shaft. It is parallel-sided for most 
of its length but for its posterior quarter tapers to a point and is free from attachment 
to the shaft. Anteriorly, it is bluntly pointed. Measurements of the baculum (in 
millimetres) : length of shaft 13-2 and length of lamina, 6-6. 

1 The surfaces of the baculum are described here according to the usage of Pocock (1923). The lamina 
is attached to that part of the shaft which in the extended penis is nearest to the belly, i.e. its upper 
or dorsal surface. The baculum, however, occurs in the distal portion of the penis which in the 
unextended condition is recurved to point posteriorly beneath the proximal portion of the penis. In 
this condition the lamina is ventrally placed in relation to the shaft. 



2 6o A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 

It has been possible to examine the bacula of specimens representative of the 
following species of Callosciurus : finlaysoni, erythraeus, flavimanus, notatus and 
prevosti. Of these, the first three have the lamina attached to the central part of 
the shaft and not extending to the tip. The lamina is pointed anteriorly, is widest 
at a point some three-quarters of its length from the tip and tapers to a posterior 
point. The tip of the shaft is furnished with two lateral barbs and with a third, 
usually larger, dorsal barb. This structure is illustrated by Didier (1952). The 
bacula of Callosciurus notatus and Callosciurus prevosti, however, resemble that of 
Glyphotes simus in having the lamina, which is the same shape as in the preceding 
species, inserted immediately posterior to the tip of the shaft. Among other genera, 
the baculum of Glyphotes appears nearest to that of Menetes, but the lamina in this 
genus, although attached near the tip of the shaft, is very small and there are no 
discernible barbs. 

The skull (Plates 7 (a), 7 (b), 8 (a)) is globular, with an exceptionally short, broad 
and deep rostrum. The junction of the nasals, premaxillae and maxillae forms an 
almost straight line joining the anterior edges of the anterior zygomatic roots. The 
nasals are short, broad and nearly parallel-sided, while the nasal aperture is flattened 
dorsally. The interorbital region is broad while the postorbital processes of the 
frontal are reduced and lie directly above the anterior edge of the posterior zygomatic 
root. The braincase is globular and not especially inflated. The orbit is more 
circular than in Callosciurus and is not placed especially far back, its anterior rim 
lying directly above p 4 , with the lacrimal lying above p 4 and m 1 . The zygomatic 
plate is weakly ridged and is slightly oblique, more vertical than in Callosciurus, 
its orbital edge ascending above the posterior face of p 4 . The zygomatic arch is 
moderately strong with a small jugal process lying at the lowest part of the arch. 
The posterior zygomatic root is formed as in Callosciurus, the orbital surfaces of the 
alisphenoid and squamosal only slightly ridged and not forming a shelf as in 
N annosciurus . The postorbital process of the jugal is rounded posteriorly as in 
Callosciurus. The anteorbital foramen is large, that part of the zygomatic plate 
lying over the infraorbital canal slightly reduced and narrow. The anterior palatine 
foramina are large and extend posteriorly to the maxillary suture. The palate is 
wide and terminates just beyond m 3 , with no pronounced posterior palatal spicule. 
The maxillary tooth rows are parallel for much of their length but converge slightly 
posteriorly while the mesopterygoid fossa is moderate and the pterygoids not 
excessively developed. A narrow ectopterygoid is present. The bullae are not 
inflated. The mandible is weak with a minute coronoid process, lower in height 
than the condylar process, which itself is slender and much elongated posteriorly. 
The angular process is narrow and unthickened. 

The incisors (Plate 8 (b)) are orthodont and very much specialized. The upper 
incisors are broadened but not thickened antero-posteriorly, their surfaces without 
grooves and with the anterior surface strongly convex. The outer lateral faces are 
strongly concave, the inner faces correspondingly convex so that for their terminal 
millimetre the teeth diverge from each other. The lower incisors are broadened, 
their anterior faces convex (not concave as stated by Thomas) and are not thickened 
antero-posteriorly. The lateral faces are similar to those of the upper incisors, the 



A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 261 

teeth similarly divergent at the tips. The lower incisors are worn into broad, flat 
chisel-shaped blades, the inner corners more worn than the outer, so that viewed 
from the front the teeth present a shallow V-shape. That these peculiarities are 
not artifacts caused during skull cleaning is proved by their presence in the spirit 
specimen examined. The cheek teeth are of normal Callosciurine pattern with p 3 
not especially reduced and the upper molars not excessively ridged. In the mandible, 
p 4 is slightly smaller than the lower molars and m 3 is not elongated, while the antero- 
internal cusps of the lower teeth are high as in Callosciurus. 

The following measurements (in millimetres) are those of an adult male (B.M. 
58.446) with measurements of the type specimen in parentheses : the skull of the 
Raffles Museum specimen is too fragmentary for measurement, but some measure- 
ments obtained from this skull and of Sarawak Museum 55.11 appear in Chasen & 
Kloss (1927). Total length of skull 29-1 ( — ), condylobasal length 25-9 ( — ), 
occipitonasal length 28-9 ( — ), basal length 23-5 ( — ), zygomatic breadth 19-0 ( — ), 
braincase breadth 16-4 (16-5), nasals, length x breadth 6-6 X 5*2 (67 X 5-3), 
diastema 6-o (57), palatal length 13-2 (12-5), height of muzzle behind incisors 6-3 
(6-i), breadth of muzzle over roots of incisors 7-5 (y-8), bulla, length x breadth 
5*5 X 3-3 (5'0 X 3-0), maxillary tooth row 4-8 (4-8), i 1 — i 1 (tips) 3-4 (3-2), length 
X breadth p 3 07 X i-i (o-6 X 07), length X breadth p 4 i-o X 1-2 (0-9 X 1-3), 
length x breadth m 1 1-1x1-3 (1-1x1-3), length X breadth m 2 1-1x1-4 
(1-2 X 1-4), length x breadth m 3 i-o X 1-2 (i-o X 1-2), orbital length 9-6 (9-5), 
length from lacrimal notch to tip of nasals 10-5 (9*6). 

Apart from its specialized incisors and greatly shortened muzzle, the skull of 
Glyphotes is similar to that of Callosciurus notatus, although, of course, much smaller. 
Certain cranial characters, however, such as the slight backward deflection of the 
components of the orbit, the obliquity of the zygomatic plate, which although more 
nearly vertical than in Callosciurus is more oblique than in N annosciurus , the 
reduction of the post orbital processes and their backward displacement, the slight 
backward displacement of the lacrimal, a more circular orbitotemporal fossa and 
the reduction of the coronoid processes tend towards N annosciurus , Prosciurillus, 
Sciurillus and Myosciurus. The structure of the baculum, the dentition and the 
major cranial characters, however, associate Glyphotes indubitably with 
Callosciurus. 

The genus Glyphotes was placed by Ellerman (1940) in his " Lariscus " section of 
Sciuridae, a heterogeneous group containing Lariscus, Menetes, Rheithrosciurus , 
Rhinosciurus and Hyosciurus (an assemblage which Ellerman admits is not a natural 
group) the principal criterion being that these genera are much specialized and 
distinct from Sciurus. Simpson (1945), following Pocock (1923), places Glyphotes 
in the Callosciurini or Oriental tree squirrels. The cranial, dental and more 
especially the bacular characters of Glyphotes however, associate the genus more 
closely with Callosciurus than was thought by Ellerman and it seems best regarded 
as an offshoot of the ancestral stem of this widespread Oriental genus. The 
remarkable resemblance in colour pattern to Callosciurus notatus appears to be the 
result of parallelism : this pattern, with slight variation, reappears in Callosciurus 
nigrovittatus. 



262 A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 

Widely divergent views exist on the classification of the dwarf Sciuridae of 
Borneo, Celebes, West Africa and South America. Earlier authors, i.e. Forsyth 
Major (1893), Thomas (1914, 1914a) and Miller & Gidley (1918) unite the dwarf 
squirrels in a subfamily, Nannosciurinae, based on certain dental and cranial 
characters common to each. A less extreme view is taken by Ellerman (1940) who 
places Nannosciurus, Sciurillus and Myosciurus in a Nannosciurus section of 
Sciuridae. By inference, the genus Prosciurillus , erected by Ellerman (1947) for 
Sciurus murinus Miiller & Schlegel, would be placed in this section by Ellerman. 
Others, i.e. Pocock (1923) deny any close relation, mainly from the study of the 
penial characters, and suggest that similarities between these genera are the result 
of convergence, a view apparently adopted by Simpson (1945), who places Nanno- 
sciurus (and by inference Prosciurillus), Sciurillus and Myosciurus with the 
Callosciurini, Sciurini and Funambulini respectively. An important but limited 
contribution is made by Anthony & Tate (1935), who compare Nannosciurus, 
Sciurillus and Sciurus and conclude that a close relation must be presumed to 
exist between Sciurillus and Nannosciurus, despite differences in their bacula. 

The skulls and bacula of Callosciurus, Glyphotes, Microsciurus, Prosciurillus, 
Sciurillus, Myosciurus and Nannosciurus have been compared, examining particu- 
larly those characters shared by the four latter, more extreme genera. Dental 
characters have not been taken into account : all but the last two genera have 
molars with the normal Sciurine ridge and depression pattern. Although suppressed 
in Myosciurus and Nannosciurus, as pointed out by Ellerman (1940) it is occasionally 
possible even so to trace the pattern in some skulls. Suppression of the pattern is 
much less evident in Prosciurillus and Sciurillus. Attention has been directed 
towards characters likely to be modified by the small size and specializations (when 
compared with Callosciurus or Sciurus) of the dwarf genera. The results are 
presented in Table I, itself extended and adapted from the tabular comparison of 
Anthony & Tate (1935). 

Moore (1958) has extended Prosciurillus as defined by Ellerman (1947) to include 
Callosciurus leucomus, a species with a shortened orbit and which Ellerman (1940) 
points out is " almost transitional" to the Nannosciurus-Sciurillus type of skull. 
There is much to commend this action : leucomus differs considerably from 
Callosciurus, sensu stricto, in the form of its orbit, in the posterior displacement of 
its post orbital processes and in the obliquity of its zygomatic plate. Ellerman 
(1949), while including leucomus within Callosciurus, in fact suggested that it might 
warrant subgeneric rank within that genus. Moore, however, does not indicate 
that at the same time leucomus differs somewhat from murinus, the type species of 
Prosciurillus. It has well developed postorbital processes (like those of Callosciurus) 
which are not displaced posteriorly as far as those of murinus : its orbit, while less 
elongate than that of Callosciurus is less circular than that of murinus : good series 
of skulls of both forms show that the lacrimal is less extruded into the margin of 
the orbit than in murinus : the coronoid process is less reduced : the articulatory 
area of the jugal and squamosal is not reduced : the zygomatic plate is more oblique 
than in murinus and the posterior edge of the nasals is less advanced from a line 
joining the anterior zygomatic roots. It is clearly transitional between Callosciurus 



A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 263 



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264 A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 

and Prosciurillus murinus and since its differences from the latter species are mainly 
small differences of degree it seems best included in that genus as denned by Moore. 
Furthermore, Prosciurillus abstrusus, described as new by Moore (1958) seems 
from the description to be intermediate in some respects between murinus and 
leucomus. Microsciurus has been included in Table I since, like Prosciurillus, it 
is a transitional genus between the true dwarf squirrels and the larger Callosciurus 
and Sciurus. Microsciurus similis, transferred to Sciurillus by Ellerman (1949), is 
considerably less specialized than that genus and for the purposes of the present 
paper is retained within Microsciurus. 

Gradual divergence from the normal condition exemplified by Callosciurus notatus 
appears in a number of characters. Some of the characters examined are at least 
in part the direct result of the shortening of the base of the skull and its arcuate 
deflection. Such characters are the circular form of the orbit, the rearward dis- 
placement of the postorbital process, the separation of the frontal process of the 
premaxilla from the lacrimal by a projection of the maxilla, the extrusion of the 
lacrimal into the orbital margin, the forward extension of the suborbit, a tendency 
for the incisors to become strongly proodont, a lessening of the obliquity of the 
zygomatic plate and alterations in the form of the nasals. Others appear related 
to this cause less for mechanical reasons but through alterations in muscle 
attachments i.e. the development of alisphenoid ridges, increases in the relative 
area of the maxillary insertion of the masseter and alterations in the articulatory 
parts of the mandible. Yet others appear adventitious, such as the presence or 
absence of a post-palatal spine, the form of the jugal process and the presence or 
absence and degree of development of the ectopterygoids. 

Sciurillus, Myosciurus and Nannosciurus are seen to be the most extreme genera. 
Of these, Myosciurus and Nannosciurus, the smallest genera, are the most divergent. 
Sciurillus occupies a position intermediate between Microsciurus, Prosciurillus and 
the extreme genera. Myosciurus and Sciurillus are further specialized in the form 
of the zygomatic plate, which in Myosciurus is greatly reduced and in Sciurillus is 
grooved above the anteorbital foramen. This aperture in Prosciurillus, Sciurillus 
and Nannosciurus lies well in advance of the anterior end of the maxillary tooth 
row. Prosciurillus and Microsciurus are transitional between Glyphotes and the 
more extreme genera and display many of their characters in lesser degree. The 
skull of Microsciurus closely resembles that of Prosciurillus, both genera displaying 
the characters considered here at about the same level of development. This 
factor no doubt led Ellerman (1940) to associate Prosciurillus murinus with 
Sciurillus and later (1949) to transfer Microsciurus similis to Sciurillus. 
Both genera, however, are less specialized than Sciurillus : Microsciurus similis 
(transitional to Sciurus) in fact stands further from Sciurillus pusillus than does 
Prosciurillus leucomus (transitional to Callosciurus) to Prosciurillus murinus. 
Glyphotes connects Prosciurillus and Callosciurus in the development of its skull 
while Microsciurus, considerably more advanced in the development of the characters 
considered, provides a more remote link between Sciurillus and Sciurus. Such 
bacula as have been examined suggest no relation between the genera. That of 
Glyphotes is typically Callosciurine, while the baculum of Sciurillus is said by 



A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 265 

Anthony & Tate (1935) to resemble that of Sciurus (Sciurus niger). The baculum 
of Myosciurus, which stands somewhat apart from other Sciurid bacula, resembles 
most closely in its small size and terminally thickened rod-like form the bacula of 
the African tree squirrels. The hook-shaped, laminated baculum of Nannosciurus 
is nearest to the " Tomeutes " type of Thomas (1915). No regular pattern is 
displayed by the adventitious characters. The form of the post orbital process of 
the jugal is variable, but its position on the arch of the zygoma appears related to 
the shortening of the orbit. It is on or near the lowest part of the arch in 
Callosciurus, Glyphotes, Microsciurus and Prosciurillus . In Myosciurus and 
Nannosciurus it is pushed forward to a position on the anterior ascending part of 
the arch, while Sciurillus displays an intermediate condition. 

There appears to be no intermediate genus comparable with Glyphotes or 
Microsciurus among the African tree squirrels. Reduction of size and relative 
shortening of the rostrum, however, occur within the genus Paraxerus, of which 
the smaller members, on account of certain external and dental characters, are 
usually considered a subgenus, Tamiscus. The larger species of Paraxerus (the 
cepapi, flavivittis and palliatus groups of Ellerman (1940)) have skulls similar to 
Callosciurus in many respects and show no trace of the specializations of dwarf 
genera. Some of these characters appear among the members of the subgenus 
Tamiscus, all of which are smaller and have a relatively shorter rostrum. The 
orbitotemporal fossa is shortened and less elongated than in the larger species and 
the postorbital processes are displaced posteriorly to lie slightly in advance of 
(boehrni, emini, vulcanorum, antoniae) or above (alexandri) the anterior face of the 
posterior zygomatic root. There is a very slight ridge on the alisphenoid and the 
upper incisors are more proodont than in the larger species. The mandible, however, 
has remained unspecialized and there is no lessening of the obliquity of the zygomatic 
plate. 

The grouping of the Celebean genus Prosciurillus, the South American genus 
Sciurillus, the West African genus Myosciurus and the Bornean genus Nannosciurus 
in one subfamily is unconvincing on zoogeographical grounds. All are highly 
specialized genera, widely divergent from Sciurus or Callosciurus, sharing in common 
a number of cranial characters which appear mostly to be related to the common 
factor of greatly reduced size. Prosciurillus, the largest, is a " giant " among 
these pygmy genera, and is the least divergent from Callosciurus. They diverge 
considerably among themselves. Myosciurus is particularly aberrant, and so far as 
comparison has been possible, their bacula have been found to differ widely, usually 
resembling in each case the bacula of the more normal tree squirrels of the region 
in which the dwarf is found. Study of the " semi-dwarf " Glyphotes, Microsciurus 
and Prosciurillus reveals that the characters appearing in extreme form among the 
most divergent genera are found in them in lesser degree and that these intermediate 
genera connect the " Nannosciurinae " to Callosciurus and Sciurus. The characters 
show difference in degree within the genus : Prosciurillus murinus ranks closely 
to the more extreme Sciurillus but the larger Prosciurillus leucomus displays con- 
siderable affinity with Callosciurus and, in the absence of the smaller species, would 
possibly warrant subgeneric rank within Callosciurus. The evidence therefore 



266 A BORNEAN PYGMY SQUIRREL, GLYPHOTES SIMUS THOMAS 

suggests that rather than forming a unit with biological reality, the dwarf squirrels 
of Celebes, Borneo, West Africa and South America have evolved independently. 
The grouping by Simpson (1945) of these squirrels with the tree squirrels of their 
respective regions appears a more accurate representation of their relationships 
than placing them in an artificial group which ignores the genera connecting them 
to the other members of the Sciuridae. 

REFERENCES 

Banks, E. 1931. A popular account of the mammals of Borneo. /. Malay Br. Roy. As. 
Soc. 9, 2 : 1-139, pis. 11-19, map. 

Chasen, F. N. & Kloss, C. B. 1927. A rare Bornean Squirrel, Glyphotes simus Thomas. 
Ibid. 5 : 349-35°- 

Didier, R. 1952. Etude systematique de l'os penien des mammiferes. Rongeurs. Mam- 
malia, 16 : 7-23. 

Ellerman, J. R. 1940. The Families and Genera of Living Rodents. Vol. 1. London : 
British Museum (Natural History). 

1947- Notes on some Asiatic Rodents in the British Museum. Proc. zool. Soc. Lond. 

117 : 259-271. 

1949. The Families and Genera of Living Rodents. Vol. 3, pt. 1. London : British 

Museum (Natural History). 

Forsyth-Major, C. J. 1893. On some Miocene squirrels, with remarks on the dentition and 

classification of the Sciuridae. Proc. zool. Soc. Lond. 179-215, pis. 8-1 1. 
Miller, G. S. & Gidley, J. W. 1918. Synopsis of the supergeneric groups of Rodents. /. 

Wash. Acad. Sci. 8 : 431-448. 
Moore, J. C. 1958. A new species and a redefinition of the squirrel Genus Prosciurillus of 

Celebes. Amer. Mus. Novit. No. 1890 : 1-5. 
Pocock, R. I. 1923. The classification of the Sciuridae. Proc. zool. Soc. Lond. 209-246, 

figs. 18-29. 
Simpson, G. G. 1945. The Principles of classification and a classification of Mammals. Bull. 

Amer. Mus. nat. Hist. 85 : i-xvi, 1-350. 
Thomas, O. 1914. Proc. zool. Soc. Lond. Abstract. No. 133 : 36. 

1914a. On a remarkable case of affinity between animals inhabiting Guiana, West 

Africa, and the Malay Archipelago. Proc. zool. Soc. Lond. 415-417. 

1915. The penis-bone, or " Baculum ", as a guide to the classification of certain squirrels. 

Ann. Mag. nat. Hist. 15 : 383-387. 



PLATE 7 

(a) Glyphotes simus. Dorsal aspect skull and ventral aspect mandible, x 3*5. 

(b) Glyphotes simus. Ventral aspect skull and dorsal aspect mandible, x 3*5. 

PLATE 8 

(a) Glyphotes simus. Lateral aspect skull and mandible, x 2*5. 

(b) Glyphotes simus. Frontal aspect incisor teeth, x 3. 

(c) Glyphotes simus. Dorsal aspect baculum. x 5. 

(d) Glyphotes simus. Lateral aspect baculum. x 5. 

(e) Glyphotes simus. Dorsal aspect tip of baculum. x 12. 
(/) Glyphotes simus. Lateral aspect tip of baculum. x 12. 



Bull. B.M. (N.H.) Zoo!. 5, 9 



PLATE 7 





Plate 7(a) 



!;.#: 





Plate 7 (ft). 



Bull. B.M. (N.H.) Zoo!. 5, 9 



PLATE 8 




**M 





Plate 8(a). 




Plate 8(b). 




Plate 8(e). 




Plate 8(d). 



Plate S(/). 



PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 

BARTHOLOMEW PRESS. DORKING 



0* 



REVISION OF THE WORLD 
SPECIES OF APLYSIA 

(GASTROPODA, OPISTHOBRANCHIA) 



N. B. EALES 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 5 No. 10 

LONDON : i960 



REVISION OF THE WORLD SPECIES 

OF APLYSIA 
(GASTROPODA, OPISTHOBRANCHIA) 



BY 

N. B. EALES, D.Sc. 

Zoology Department, University of Reading 




Pp. 267-404 ; Frontispiece ; 51 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 5 No. 10 

LONDON: i960 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 5 No. 10 of the Zoological series. 



© Trustees of the British Museum, i960 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued January, i960 Price Forty -five Shillings 



REVISION OF THE WORLD SPECIES 

OF APLYSIA 
(GASTROPODA, OPISTHOBRANCHIA) 

By N. B. EALES, D.Sc. 

CONTENTS 

Page 
Synopsis and List of New Species ....... 269 

1. Introduction .......... 269 

(a) Definition of the subfamily and genus . . . . .269 

(b) Subgenera and list of species . . . . . . .271 

(c) History of the genus . . . . . . . .274 

(d) Practical notes . . . . . . . . . 277 

(e) Criteria employed in description . . . . . .286 

2. Descriptions of the Species ....... 287 

3. Geographical Distribution ........ 377 

4. Conclusions ........... 380 

5. Acknowledgments ......... 382 

6. Species Represented in Museum Collections .... 383 

7. Reference Letters ......... 392 

8. Literature ........... 392 

9. Index to Specific Names ........ 401 

SYNOPSIS 

A review of the genus Aplysia and of the geographical distribution of its members has been 
made, based on the collections of the British Museum (Natural History) with comparative 
material from other national museums of Natural History, Marine Biological Stations and private 
collectors all over the world. The genus can be divided into five subgenera : Pruvotaplysia, 
the most primitive, with two species ; Neaplysia, North American, with probably only one 
species ; Varria nov., with more than twenty species ; Aplysia, with six species ; and Phyco- 
phila, of the Pacific, with one, possibly two species. 

Full descriptions of the species are given, and the following new species are described : 
cronullae, dura, gracilis, rehderi, reticulata, as well as a new variety, delli of A. nigra. Their 
affinities with one another and with other subfamilies of the Aplysiidae are discussed. 

1. INTRODUCTION 

(a) Definition of the Subfamily and Genus 

The Tectibranch family Aplysiidae contains four subfamilies : 

Aplysiinae : Syphonota and Aplysia. 
Dolabellinae : Dolabella. 

Dolabriferinae : Dolabrifera, Petalifera, Phyllaplysia. 
Notarchinae : Notarchus, Stylocheilus , Barnardaclesia, Bursatella. 
zool. 5, 10. 13 



270 REVISION OF THE WORLD SPECIES OF APLYSIA 

The Aplysiinae are mostly large molluscs, with humped body, the skin usually 
with reticulate black markings, but without warts or villi. There are no separate 
oral lobes. The cephalic tentacles are broad and folded, the rhinophores slit like 
hare's ears. The foot has a short or moderately elongated tail. The parapodia 
are well developed, symmetrical, mobile, upstanding, separated anteriorly, free or 
joined posteriorly, typically natatory. The mantle encloses the shell, with or without 
a mantle foramen. Sub-pallial glands secrete either a purple or a white fluid. The 
anal siphon is usually tubular and erect, the opaline gland simple or compound. 
The genital aperture is inside the dorsal slit, anterior to the ctenidium. Shell narrow 
to broad, flexible, with reduced calcareous layer and apex, with an anal sinus on 
the right posterior border. Jaws large, radula with wide, denticulate rhachidian 
tooth and bi-serrate laterals. Caecum with a typhlosole present. Penis unarmed, 
penis sheath rarely armed, no penial collar. Nerve ring with three pairs of ganglia, 
pleuro visceral cords long, supra-intestinal and visceral ganglia adjacent, forming 
a pair or fused. There are two genera, Syphonota and Aplysia. 

SYPHONOTA Adams, 1854 1 

Head broad, with rhinophores set close together and far back between the anterior 
ends of the parapodia. Foot broad. Parapodia free posteriorly, the backwardly 
directed anal siphon projecting through the gap between them, the shell and mantle 
tilted towards the animal's left side. Purple glands present. Opaline gland simple, 
multiporous. Genital aperture almost mid-dorsal, not covered by the mantle flap. 
Caecum not reaching the surface of the digestive gland. 

Type species Siphonotns geographica Adams & Reeve, 1850. Cir cum tropical. 
Probably only one species. Baba (1955, pi. 1) gave a good coloured figure of this 
species. 

APLYSIA L., 1767 

Body soft in most species, elongated but strongly contractile, assuming the 
" sitting hare " position. The head is more shapely than in Syphonota, with erect 
rhinophores not far distant from the cephalic tentacles, so that a neck region is 
present between the head and the parapodial attachment. The foot varies from 
narrow to medium or broad, with a relatively short tail. The upstanding parapodia 
are mobile and usually natatory, joined high up in some species, shutting in the 
mantle cavity posteriorly ; meeting or joining low down on the foot in others, so 
that the mantle cavity, as in Syphonota, is open at both ends. Anal siphon erect. 
Secretion from the mantle glands purple except in the subgenus Aplysia where it 
is white or black. Opaline gland simple or compound, multiporous or uniporous 
respectively. Penis varying from short and broad to long and filiform, penis sheath 
smooth distally except in subgen. Aplysia where it is lined with warts bearing spines. 
Shell ovate or rounded, horny, with an imperfect calcareous lining ; in the subgenus 
Neaplysia the apex of the shell is flattened and rectangular, in all others recurved 
or hooked. Radula large, multiserial, all the teeth denticulate in most species 

1 To replace Siphonotus Adams & Reeve, 1850 (nee Brandt, 1837). 



REVISION OF THE WORLD SPECIES OF APLYSIA 271 

except for a few vestigial teeth at the end of the row. Caecum appearing on the surface 
of the digestive gland. Cerebral ganglia distinct or fused. Parietal (supra-intestinal) 
and visceral ganglia free or fused. 

Numerous species, most of which are tropical ; others live in the temperate zone 
up to high latitudes. 

Type species A. depilans Gmelin. European. 

(b) Subgenera 

The genus Aplysia can be divided into five subgenera : 

1. Pruvotaplysia Engel, 1936, type Aplysia parvula Guilding in Morch, 1863, 
a primitive subgenus with two species. 

2. Neaplysia Cooper, 1863, type and only species A. (N.) californica Cooper. 

3. Varria 1 no v., type Aplysia dactylomela Rang, 1828. Twenty-five species are 
here recognized. 

4. Aplysia Linnaeus, 1767 (as Laplysia, probably a printer's error), type A. 
depilans Gmelin, 1791A Six species, on one of which, A. Juliana Quoy & Gaimard, 
was founded the subgenus Tullia by Mme. Pruvot-Fol, who did not realize that its 
peculiarities were shared by the type species of the genus. 

5. Phycophila A. Adams, 1861, type Placobranchus euchlora M. E. Gray, 1850 
(ex Adams unpublished). This subgenus includes oceanic floating Aplysias known 
in the Pacific Ocean and belonging to one or possibly two species. 

Characters of the Subgenera 

1. Pruvotaplysia. The foot is narrow, with a pointed tail ; the parapodia are joined 
high up posteriorly, closing in the mantle cavity behind. There is a flat, unrayed 
mantle aperture over the strongly concave shell, which usually has remnants of 
a spiral at its apex. The mantle glands secrete purple, the opaline gland is simple 
and multiporous, the radula small, with not more than 40 rows and fewer than 
25 laterals on each side in a row. The ganglia of the nerve ring and of the visceral 
group (supra-intestinal and infra-intestinal plus visceral) are rounded and distinct. 
All these features can be regarded as primitive. There are two species, parvula 
the type and punctata, the former circumtropical, the latter in the north temperate 
and arctic Atlantic areas of the Old World. 

2. Neaplysia, with one species, californica. Giant forms, with strongly mottled 
skin. Foot broad, but with a long slender tail. Parapodia joined posteriorly low 
down. Mantle aperture minute or closed. Purple secretion from the mantle glands. 
Opaline gland large, compound, uniporous. The only peculiarity of this subgenus 
is the shell, which has a flattened calcareous apex, instead of a reduced spiral or 
hook. This rectangular flattening was described by Cooper (1863) as an accessory 
plate, but is part of the shell itself. In all other features the affinities are with 
subgenus Varria. The species is confined to the eastern North Pacific area. 

3. Varria. This group contains about seven-tenths of the known species, which 

1 Varria from Varro, the name of a famous Roman scholar and writer. 

2 Opinion 200 of the International Commission on Zoological Nomenclature. See p. 396, 



272 REVISION OF THE WORLD SPECIES OF APLYSIA 

show remarkable generalization of structure, despite a great range of size. The 
parapodia are joined low down posteriorly, exposing the mantle cavity to a through 
current, since it is open at both ends, and although the anal spout is typically erect, 
it can be directed backwards between the parapodia as in Syphonota. In a few 
species, e.g. inca, rehderi, preserved specimens exhibit a low wall not more than 10 
mm. high posteriorly, but in the living animal this probably forms a flat extension 
of the mantle floor, as it does in other species. It is quite unlike the high fusion in 
the subgenera Pruvotaplysia and Aplysia. There is always a purple secretion from 
the mantle glands. About half the members of this group have a simple multiporous 
opaline gland, the other half a compound uniporous gland, with gradations between 
them in some species, e.g. cornigera, keraudreni, pulmonica. The foot is narrow in 
the majority, of medium width in cervina, oculifera and winneba, and broad in 
dactylomela, denisoni, gigantea, keraudreni and pulmonica. The penis is filiform in 
cronullae, fasciata, keraudreni, kurodai, sowerbyi, Sydney ensis, willcoxi and winneba, 
broad and spatulate in dactylomela, denisoni, gigantea and pulmonica, but tapering 
in the remaining species. The shell varies, but it never possesses a flat, plate-like 
apex, and is usually rather shallow. The radula is well developed, with many rows 
(up to about 80) and numerous laterals (to approximately 60) ; the laterals are 
multidenticulate except in dactylomela and pulmonica, where the denticulations 
exhibit feeble development in young teeth but disappear later, producing long, 
smooth-edged cusps, usually with a single stout lateral denticle. The caecum is 
straight or only slightly bent at the tip. The nerve ganglia exhibit fusion of the 
cerebral and of the visceral pair, except in sagamiana, which has small rounded 
discrete ganglia as in Pruvotaplysia. In its small size, simple multiporous opaline 
gland and narrow foot, sagamiana resembles parvula, but in its small rayed mantle 
aperture, rather flat shell without spiral apex and low junction of the parapodia 
posteriorly its affinities are with Varria. The distribution is in all oceans except 
the Arctic and Antarctic regions. Type A. dactylomela. 

4. The subgenus Aplysia includes the type species of the genus, A. depilans. The 
body is low and flat, bulky, with a broad head. There is a broad foot, capable in 
most species of producing posteriorly a rounded but temporary sucker. The rather 
small parapodia are joined high up posteriorly. There is a flat mantle aperture. 
Purple is not secreted, but both the mantle glands and the simple, multiporous 
opaline gland secrete a white fluid, rarely a black, which is copious and odoriferous 
in the case of the opaline gland. The radular teeth are simple, but the number of 
rows and of teeth in a row reaches the maximum for the genus. The salivary glands 
are broad and flat, the caecum spiral at the tip. The penis is stout, heavily pigmented, 
and lies in a sheath which is armed with spiny warts. Six species are known. 
Distribution circumglobal in tropics and sub-tropics. 

5. Phycophila. This includes small floating species, about which little is known. 
Only one species, euchlora, from the western Pacific area, has been described. The 
cephalic tentacles are broad, curved and pointed, the body slender, with a narrow 
foot and long, tapering tail. The parapodia are freely mobile and meet low down 
on the tail. The mantle is small, the shell sac closed. The opaline gland is small, 
simple and multiporous. The penis is filiform. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



273 



List of Recognized Species of Aplysia 











Page 


Subgenus Varria (cont.): 






Page 


Subgenus Pruvotaplysia : 


morio . ..... 328 


parvula ..... 287 


oculifera 








332 


punctata . . . . .291 


pulmonica 
rehderi 








335 

337 


Subgenus Neaplysia : 


reticulata 








34° 


calif ornica . . . . .294 


robertsi 
sagamiana 








342 
343 


Subgenus Varria : 


sowerbyi 








345 


brasiliana ..... 297 


sydneyensis . 








348 


cervina 








299 


willcoxi 








350 


comigera 








302 


winneba 








352 


cronullae 








304 




dactylomela . 








307 


Subgenus Aplysia : 


denisoni 








310 


cedrosensis . . . .354 


extraordinaria 








312 


depilans 








357 


fasciata 








315 


dura 








360 


gigantea 








3i8 


Juliana 








363 


gracilis 








320 


nigra . 








369 


inca 








321 


vaccaria 








37i 


keraudreni 








322 




kurodai 








325 


Subgenus Phycophila : 


maculata 








326 


euchlora 








• 374 



Rejected Names 

Names considered invalid or of doubtful validity because of inadequate description 
and absence of a specimen. It is possible that some of these are synonyms of known 
species, but there is not sufficient information to establish their identity. 

adamsi Pilsbry, 1896 : 112. Borneo. 

chierchiana Mazzarelli & Zuccardi, 1892 : 13. Peru. 

fusca Tilesius, 1809-13 : China and Japan. 

laevigata Stimpson, 1855 : 378. China. 

marginata and marmorea A. Adams, 1861 : 140. China. 

punctatella Bergh, 1902 : 341. Philippines. 

sinensis Sowerby, 1869 : China. 

spuria Krauss, 1848 : 71. South Africa. 

tarda Verrill, 1901 : 26. Bermuda. 

tryonii Meinertzhagen, 1880 : 270. New Zealand. 

venosa Hutton, 1875 : 279. New Zealand. 

vexans Bergh, 1905 : 10. Amboina, East Indies. 

Siphonota elongaia Pease, i860 : 24. Hawaii. 

Siphonota grandis Pease, i860 : 23. Hawaii. 



Names given to juvenile forms whose status is doubtful : 

parva Pruvot-Fol, 1953 : 38. Morocco. According to Marcus, 1958 : 10 this 
is a young Juliana. 



274 REVISION OF THE WORLD SPECIES OF APLYSIA 

sorex Rang, 1828 : 57 (see p. 363). 

Esmia griffithsiana Leach, 1847 : 266 (see p. 292). 

vistosa Pruvot-Fol, 1953 : 37. Morocco. 

Species named from shells only and unidentifiable from the descriptions : bipartita, 
elegans and macula Turton, 1932 : South Africa. 

Name given to a probable hybrid : 

Siphonota lobiancoi Mazzarelli, 1890 : 42 (p. 315). 

(c) History of the Genus 

Aplysia has been known from ancient times, the first authentic description being 
that of Pliny in the first century, a.d. He called it Lepus marinus, the Sea Hare, 
and described three kinds, two in the Mediterranean, and one, probably a Dolabella, 
in the Indian Ocean. He gave an account of the supposed poisonous qualities of 
this harmless mollusc. Other early writers compared it with a snail (Aelian) or 
a cuttlefish (Dioscorides) . 

In 1554 Rondelet included three kinds of Lepus marinus among the fishes in 
his Be Piscibus Marinis, the first of which we now know as the Nudibranch Tethys 
leporina. He explained that it should not be confused with other fishes because 
"it is very poisonous and would be fatal to anyone who ate it ". But Gesner in 
1551 had recognized that it was not a fish and placed it among the soft-bodied 
animals. 

Redi in 1684 called it the Sea Slug and described its internal anatomy, and a 
century afterwards Bohadsch, a Bohemian fugitive living in Naples, published in 
1761 a book on marine animals, describing two kinds under the name of Lernaea. 
He studied their habits and observed the discharge of purple and milk-white secre- 
tions. The opaline gland is sometimes called by his name. He realized that his 
Lernaea is related to the land snail. 

British observers in the eighteenth century classified Aplysia amongst the Holo- 
thurians (Borlase, 1758) and the worms (Pennant, 1777). 

In 1756 Linnaeus, in the 9th edition of the Systema Naturae, used Lernea (later 
adopted by Bohadsch as Lernaea) for the Sea Hare ; in the 10th edition (1758) he 
changed it to Tethys, mentioning two species, which he called T. limacina, with the 
habitat " Oceano Australi " and T. leporina (= Lepus marinus) in the Mediterranean; 
in the 12th edition, 1767 he changed the diagnosis, applied the name Tethys to the 
animal we know now as a Nudibranch and called the Sea Hare Laplysia (probably 
a printer's error), choosing as the type L. depilans 1 ; finally in the 13th edition 
(1791) Gmelin corrected the spelling to Aplysia. 

1 The word Aplysia means " that which one cannot wash " and was used by Aristotle for a sponge 
which could not be freed from grit and sand. Linnaeus chose the name arbitrarily. Aplysia came into 
common usage for the Tectibranch, Tethys for the Nudibranch. On the question of priority Pilsbry 
changed Aplysia to Tethys, thus making Tethys a genus of the family Aplysiidae and Aplysia a genus 
of the family Tethyidae. The International Commission on Zoological Nomenclature (Op. 200, 1954, 
3, 239-266) decided that the name Aplysia should be validated for the Tectibranch notwithstanding the 
Linnaean name in the 10th edition. 



REVISION OF THE WORLD SPECIES OF APLYSIA 275 

In the nineteenth century Lamarck (1822) classified the Aplysiidae near the Bulliidae. 
Cuvier in 1803 published a memoir on the genus " Laplysia " and gave the first 
full account of its anatomy, with good figures. European and especially Mediter- 
ranean species occupied the attention of Risso (1818), delle Chiaje (1828), de Blain- 
ville (1823), Milne Edwards (1847), Vayssiere (1885) and Mazzarelli (1893), while 
the great voyages of the period brought specimens from foreign coasts in all parts 
of the world. Rang's monograph Histoire naturelle des Aplysiens, 1828 is still valuable 
for the descriptions and figures it contains. He accounts for 22 species of the genus 
Aplysia, which the modern systematist would reduce to seven or eight. He knew 
the three Mediterranean species punctata, depilans and fasciata, but did not realize 
that some species are circum tropical. In fact, this has always caused confusion 
in the genus. Local names have been given to world-wide species, thus multiplying 
the number of synonyms. Rang's species are brasiliana, dactylomela, maculata, 
keraudreni, depilans, fasciata and punctata. 

At the end of the century the great monograph of Tryon appeared, Pilsbry con- 
tributing the section on the Aplysiidae, published in 1895 and 1896. Although 
largely a compilation from many authors it is the most comprehensive work 
existing on the family. He records at least 20 authentic species under the name 
Tethys, with numerous synonyms. 

The expansion of our knowledge in the nineteenth and twentieth centuries is due 
to two groups of workers. The marine expeditions collected material from all the 
oceans, depositing it in the National Museums of the countries concerned. In some 
cases it was investigated, but often it was left unnamed. Some of these collections 
have been used by the author and provide interesting finds. For example, specimens 
of Aplysia extraordinaria from New South Wales, a species named by Miss Allan 
in 1932, have been in the Paris National Museum since 1874 and in the British 
Museum (Natural History) since 1883. Aplysia morio, described by Verrill from 
Bermudan examples in 1901, has been represented by a good specimen in the Swedish 
national museum since 1889. 

The other group of workers consists of the local collectors who name their own 
species, often without access to known species for comparison, or to the literature. 
Frequently they failed to recognize world-wide species, with the result that synonyms 
were multiplied, young individuals were sometimes given specific rank and imperfect 
descriptions may make the species unrecognizable. Still more reprehensible was 
the naming of species from the dried shell alone, for shell shape is one of the most 
variable characters in the genus. To all who undertake to name new species, the 
author would recommend a study of p. 277 which gives the characters necessary 
for the identification of a species of Aplysia. 

The seas of the world may be divided into ten areas according to the distribution 
of species of Aplysia (see Geographical Distribution, p. 377). This division is con- 
venient but not arbitrary. Each area has been investigated by one or both of the 
two groups of zoologists mentioned above. 

The three European species punctata, fasciata and depilans are perhaps best 
known, although synonyms are numerous. Work on them during the nineteenth 
century was initiated by Cuvier (1803 and 1817), continued by the compilations of 



276 REVISION OF THE WORLD SPECIES OF APLYSIA 

de Blainville (1823 and 1825) and culminated in the beautiful and exhaustive mono- 
graph by Mazzarelli in 1893. In Great Britain only one species is indigenous, as 
Garstang (1890) recognized, but occasionally the larger continental species appeared 
on the south coasts and were recorded by Couch (1870) and Hunt (1877), and were 
identified by Grigg (1949). An account of the anatomy of A. punctata was published 
by the present author in 192 1. 

On the Atlantic coasts of North America the earliest systematist was Morch, 
who in 1863 and 1875 named species from the West Indies. Thiele (1910) described, 
not very successfully, a few species from the same area, and Verrill (1901) gave the 
first account of a common, but hitherto unnamed, North American species, to which 
he gave the name Aplysia morio. Engel (1927 and 1936), in his work on the three 
species dactylomela, cervina and parvula from the West Indies, was the first to 
recognize that some species, such as dactylomela, are circum tropical. Further north 
Heilprin (1886) found and named A. willcoxi from New Jersey, and Sanford (1922) 
extended its range to New England. 

In the West African region Risbec (1931) and Mme. Pruvot-Fol (1953) have 
studied the species from Morocco to Senegal, Odhner (1932) those of the Canaries 
and the author (1957) a collection from the coast of Ghana. 

The South Atlantic has had little investigation. Bergh (1898) wrote one of his 
earliest papers on Opisthobranchs from the Plate, MacFarland (1909) and von 
Jhering (1915) described the Brazilian species, and more recently Dr. and Mrs. 
Marcus (1955) have published valuable papers on the Opisthobranchs of Brazil, two 
of which deal with Sea Hares, and have given much useful information on the living 
Aplysias to the author. 

The Indian Ocean has always been a favourite collecting area for malacologists, 
particularly the Red Sea, and the shores of India, Ceylon, the Seychelles, Mauritius 
and Madagascar. Riippell & Leuckart (1828) worked on Red Sea species, Quoy & 
Gaimard (1832), Adams & Reeve (1850), Martens (1880) and Eales (1944) on the 
island faunas, while for South Africa we have the publications of Burne (1906), 
Bergh (1908), O'Donoghue (1929) and Macnae (1955 and 1957). This region therefore 
has been well studied. Further east, China, Japan and the East Indies provided 
a happy hunting ground for the zoologist. Arthur Adams (1861) listed species from 
China and Japan, of which unfortunately there are no recorded specimens extant. 
Bergh's massive volumes on the mollusca of the Danish Expedition to Siam (1902), 
of Semper's Reisen im Archipel der Philippinen (1902 and 1908) and of the Siboga 
Expedition (1905) describe many, but not always authentic, species. Within the 
last few years Baba (1936, 1937, 1949 and 1952) has made excellent contributions 
to our knowledge of the fauna of Japan and Formosa. 

The South Pacific, including Australia, New Zealand and Polynesia has been 
fairly adequately investigated by Angas (1867, 1877), E. A. Smith (" Alert ", 1884), 
Hedley (1916, 1918, 1923), O'Donoghue (1924, Abrolhos Isl.) and Allan (1932), 
the last of whom named from living material collected at Sydney the species A. 
extraordinaria, specimens of which had been in European museums for more than 
half a century. In New Zealand Meinertzhagen (1880) described that debatable 
species A. tryonii, the type of which has been lost. Hutton (1875 an( l x 882) described 



REVISION OF THE WORLD SPECIES OF APLYSIA 277 

local species and Suter (1913) compiled a useful Manual of the New Zealand Mollusca. 
In Polynesia Eliot (1899), so well known for his studies of Nudibranchs, recorded 
some tectibranchs from Samoa, notably nigrocincta (= parvula) and benedicti 
(= dactylomela) and Gould (1852) contributed A. pulmonica from Hawaii. I agree 
with Bergh and Mme. Pruvot-Fol that Pease's species should be ignored, being 
based neither on adequate description nor on illustration. They are unrecognizable 
and no types exist. Ostergaard (1955) gave good figures of the species he considered 
to be Pease's Syphonota elongata, which may be A. parvula. 

The Caliiornian area produces some of the giants of the genus, but is not rich in 
species. Cooper (1863) described A. calif ornica, which according to Macginitie 
attains a weight of 5 lb. 12 oz. MacFarland (1924) recorded A. parvula from this 
region. From the Gulf of California Bartsch & Rehder (1939) named A. cedrosensis, 
which was later found to be related to A. depilans, and Winkler (1955) described 
a black Aplysia without a sucker, which he named A. vaccaria. 

The South American coasts from Columbia to Chile have been little worked, and 
few museums have specimens from this area. Lesson (1830), in collaboration with 
Rang, recorded A. lessoni 1 from Peru, d'Orbigny (1837) g ay e an account of several 
species, including three from Peru, which he called A. nigra, A. rangiana and A. inca, 
the second probably a variety of Juliana, and the third, the type of which is in the 
Paris National Museum, an authentic species. Mazzarelli & Zuccardi's A . chierchiana, 
1889 is a doubtful species. 

Although small collections of species of Aplysia have been described, no one has 
hitherto attempted a world survey of the genus, based on adequate structural data. 
The fact that some species are circumglobal and others purely local makes such 
a survey both difficult and necessary. 



(d) Practical Notes 

Species of Aplysia can be recognized only by a combination of characters, the 
least reliable of which are colour and shape. Colour varies with food, age and habitat, 
and may fade altogether with preservation. Shape and size change greatly unless 
special precautions are taken during fixation. 

It would help in identification, however, if collectors would record, in 
the living animal, the colour and markings, the nature of the mantle gland 
exudate (purple or white), the presence or absence of a sucker on the posterior 
end of the foot and the length of tail relative to the body. 

A species of Aplysia cannot be said to be adequately described unless the following 
features have been investigated : 

Size, appearance, mode of contraction, colour and markings, head, neck, foot, 
penis and penis sheath, parapodia, mantle and mantle aperture, anal siphon, mantle 
glands, opaline gland, shell, jaws, radula, caecum, nervous system. 

In the descriptions which follow the features investigated for the diagnosis of 
species will be arranged in the following order. 

1 Probably keraudreni. 



2 7 8 



REVISION OF THE WORLD SPECIES OF APLYSIA 



1. External form (Text-fig. i). Size, length, breadth and height. Shape and method 
of contraction. Nature of the skin, soft or leathery. Skin pigment and markings. 
Any known features of the living animal, e.g. swimming powers, curling up in a 
ball (Text-fig. 440). 

2. Head and neck. Relative size. Cephalic tentacles and oral extensions. Eyes. 
Rhinophores (Text-fig. 1). 

mant. 



pap 



an.s 



an. 




Fig. 



Aplysia dactylomela. 



ceph.t. 



Lateral view to show main parts of its external structure, x 
For reference letters see p. 392. 



3. Foot (Text-fig. 3). Soft or tough. Size, particularly relative breadth. Tail. 
Penis and penis sheath (Text-fig. 5, b, c). Parapodia, size, height, freedom and 
flexibility, anterior and posterior extent, relation to dorsal slit. 

4. Mantle. Relative size. Shell foramen, anal siphon, overhang, mantle glands, 
and nature of exudate (Text-fig. 4) . 

5. Mantle cavity. Relative size. Ctenidium, opaline gland and aperture, genital 
aperture (Text-figs. 4, 50). 

6. Shell. Size, shape, quality, horny and calcareous layers, apex, " accessory 
plate ", anal sinus (Text-fig. 6). 

7. Jaws. Size and shape. Jaw rods (Text-fig. 7, e,f). 

8. Radula. Size and shape, rows of teeth and formula, denticulations and special 
features (Text-fig. 7, a-d). 

9. Caecum. Size and shape. Other features of digestive system (Fig. 8, b-d). 

10. Nervous system. Ganglia of the nerve ring. Visceral (supra-intestinal and 
visceral) ganglia (Text-fig. 8a). 

Using Aplysia dactylomela and certain other species as examples, each feature 
used in the examination of a species is figured. 



REVISION OF THE WORLD SPECIES OF APLYSIA 279 

Colour. Colour means colour pattern as well as shade of colour. The colour is 
composed of at least two ingredients : 

(a) Transparent colour, e.g. green, yellow, red, violet, rich brown, blue ; this 
dissolves out or fades quickly in preservative, especially in alcohol. 

(b) Granular colour, usually white, black or brown, which fades or dissolves 
gradually and may last for many years. 

The soluble colour is said to be extracted from the weeds on which Aplysia feeds. 
Thus a young A . punctata, feeding on the red weed Delesseria and living in shallow 
water, is rose-red, coloured exactly like the weed — the A. rosea of Rathke. As it 
grows older the mollusc migrates shorewards to the brown weed zone and feeds 
chiefly on Laminaria, changing in colour to the rich olive brown of that weed. 
Finally the animal moves to the inter-tidal zone, and feeds on Fucus, darkening 
in colour as it does so. See Winkler 1959a. 




Fig. 2. Aplysia dactylomela. Two rings and reticulations, 
from a West African specimen, x f . 



The basic colour may be modified by the formation of spots or flecks, sometimes 
in the form of light centres to regular or irregular rings of black granular pigment, 
as in oculifera and dactylomela (Text-fig. 2). More frequently the granular pigment 
consists of irregular bands and blotches of dark colour, alternating with areas of 
unpigmented or less heavily pigmented skin. In almost all species reticulations of 
granular pigment are present, and there may be white granules, probably of a 
calcareous nature, the latter occurring also on the surface of the digestive gland. 
Species of Aplysia, however, rarely show the pattern of " ocelli ", with red or 
turquoise centres, so common in the Notarchinae. 

Foot (Text-fig. 3, a-c). This may be broad, of medium width or narrow (see 
Criteria, p. 286). There is usually a rounded or pointed tail, not as long as in Stylo- 
cheikts. In the subgenus Aplysia the posterior end of the foot forms a temporary 
sucker owing to the abundant secretion of the posterior pedal mucus-forming glands. 

Penis and penis sheath (Text-fig. 5, b, c). These are derivatives of the foot. The 
former may be short and broad, pike-shaped or long and filiform. The sheath is 
composed of two parts, the proximal portion near the aperture being strongly 
muscular, with longitudinal ridges internally. The distal portion, from which the 
penis springs, is enlarged and smooth-lined in most species, but in the subgenus 
Aplysia it is greatly enlarged and its lining bears numerous rounded swellings or 
warts, each capped by curved spines, probably chitinous in nature. 



28o 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Parapodia (Text-fig. i). The parapodia are lateral upgrowths of the foot in the 
visceral region ; they vary greatly in extent and flexibility. In most species they 
are natatory, widely spaced anteriorly and meeting or joining posteriorly. When 
joined high up, as in the subgenera Pruvotaplysia and Aplysia, they shut in the mantle 
cavity behind, forming a mantle cavity pocket. When joined or meeting low on the 
tail, as in Neaplysia and Varria, the mantle cavity is open both in front and behind, 
and the floor may be extended backwards as a flat platform. The anal siphon, 
usually erect, can then be directed backwards as in the allied genus Syphonota. 



c) 





ped.su 



Fig. 3. The foot. 

(a) Aplysia dactylomela. Foot sole broad, with rounded tail. 

(b) A . Juliana. Foot broad, with anterior side wings and posterior sucker. 

(c) A. parvula. Foot narrow, with pointed tail, contracting to form a characteristic 
wavy outline laterally. All x § . 



Mantle (Text-fig. 4). The mantle in all adult Aplysias is reflected upwards over 
the shell, enclosing the latter in a shell sac. In the very young animal the shell 
is exposed, but during growth the mantle turns up over its edge, the aperture becom- 
ing progressively smaller with age. In Pruvotaplysia and Aplysia this aperture 
remains as a round or oval flat hole, contractile in Aplysia so that it may close or 
nearly close. In Neaplysia and Varria the aperture is small, papillate or absent 
altogether. Thus the condition of the mantle aperture may vary considerably 
according to the degree of contraction of the mantle aperture. 

Opaline gland (Text-figs. $a, 9). The opaline gland, or gland of Bohadsch, is 
situated on the floor of the mantle cavity postero-lateral to the common genital 
aperture. There are two main types in the genus : 

(a) The simple gland, probably primitive, in which the gland is composed of large, 
flask-shaped cells, each with its own aperture to the mantle cavity, s.g. Pruvotaplysia, 
Aplysia, etc. 

(b) The compound gland, in which the gland cells combine to discharge by a single 
duct into the mantle cavity, the gland hanging from the floor into the haemocoele 



REVISION OF THE WORLD SPECIES OF APLYSIA 281 

like a bunch of grapes on a vine. This type occurs in Neaplysia and in about half 
of the members of Varria. 



pap. 



an.s 




mant.cav. / ct. 

para. op.gLap. 

Fig. 4. Aplysia dactylomela. Lateral view of the posterior region to show the mantle 
cavity and its contents. The right parapodium has been reflected, x 1. 

a) 




op.gl. 




ret.pn 



Fig. 5. Aplysia dactylomela. 

(a) Opaline gland in longitudinal section to show the central chamber into which the gland 
cells open, and the single aperture to the mantle cavity. This is the compound 
uniporous type, x 2. 

(b) Penis. The sperm groove is indicated by a dotted line and twists spirally round 
the penis, x 2. 

(c) Penis sheath seen from the ventral side to show the arrangement of the retractor 
muscles, x f . 



There is a tendency, in species with simple glands, e.g. cornigera, keraudreni, 
oculifera, for a few of the gland vesicles to unite, while others remain separate, 
giving one aperture, usually the anterior, larger than the others, with a cluster of 
small ones. This transition type is regarded as simple and multiporous. 



ZOOL. 5, IO. 



14 



282 



REVISION OF THE WORLD SPECIES OF APLYSIA 

b 




cm. si 

apex 

Fig. 6. Aplysia dactylomela. Shell, from an African specimen 112 mm. long, x f . 

(a) Dorsal view, showing the recurved edge. 

(b) Ventral view, showing the apex. The dotted line represents the extent of the 
calcareous layer. 



c) 




Fig. 7. Aplysia dactylomela. a-d teeth from the radula. x 84. 

(a) Rhachidian and first lateral of one side. 

(b) First to fourth laterals. 

(c) Part of three rows of laterals from the middle of a side to show the straight over- 
lapping cusps. Only three of the basal plates are figured. 

(d) Four outermost laterals. 

(e) Jaw rods, x 168. 
(/) Jaw. x f. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



283 



Shell (Text-fig. 6). The shell is a variable feature in Aplysia and is usually unre- 
liable for diagnosis of the species when considered alone. This is to be expected, 
since it is enclosed in the mantle and can only be regarded as degenerate. It consists 
of a horny exterior which is delicate and flexible, covering a smaller and often 
fragmented calcareous layer. When dried the horny portion alone is left as a rule, 
and this is not representative of the shell as a whole. Sowerby's numerous species, 



^g.buc 



a) 



pl.visc: 
conn 




g.v. 



Fig. 8. Aplysia dactylomela. 

(a) Nervous system in dorsal view. The cerebral ganglia are completely fused, also the 
supra-intestinal and visceral, x 2. 

(b) A. dactylomela. Visceral mass in ventral view to show the position of the caecum, 
straight in this species. The hermaphrodite gland occupies the posterior tip of the 
visceral mass, x i\. 

(c) Caecum of A . punctata for comparison. 

(d) Caecum of A . Juliana. 



founded on the dried shell, are therefore of little value in many cases. Some species, 
e.g. Juliana, have geographic races characterized by the shape of the shell. 
The genus has three types of shell : 

(a) Shell with a persistent involuted or spiral apex, strongly concave, e.g. parvula 
(Text-fig. 116). 

(b) Shell with apex flattened to form a rectangular plate, described incorrectly 
by Cooper (1863) as an " accessory plate ", e.g. californica (Text-fig. 12, a-c). 

(c) Shell without spire or accessory plate, usually with recurved but not flattened 
apex. Most species (Text-fig. 6, a, b). 



28 4 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Radula (Text-fig. 7, a-d). The radula is well developed and consists of many 
rows of teeth, with a formula X . 1 . X, where X varies from about 15 in small species 
to more than 80 in large ones. All the laterals are basically similar, with the excep- 
tion of from three to six of the outermost teeth, which are vestigial. 

Caecum (Text-fig. 8, b-d). The caecum appears on the surface of the digestive 
gland at its tip only, or for part of its length. There are three types, a straight 



la. hcrm. d. 




1. pi. vise, conn... 



Fig. 9. Aplysia punctata. Transverse section of a young individual through the 
visceral hump, above which lie the mantle cavity, mantle and shell. The mantle 
fold is just beginning to enclose the shell. Note the simple multiporous type of opaline 
gland, x io'5. Reproduced from L.M.B.C. Memoir 24. 



caecum lying flat on the surface (dactylomela) , a caecum curved at the tip and just 
breaking surface {punctata) and a much curved, crozier-shaped or spiral caecum, 
with the tip hidden (s.g. Aplysia). 

Nervous system (Text-fig. 8a). The nerve ring exhibits differing degrees of fusion. 
In the most primitive species (Pruvotaplysia) the cerebral, pedal and pleural ganglia 
are rounded and distinct, the cerebral and pedal ganglia with commissures. In 
most species the cerebral ganglia are contiguous within a connective tissue sheath, 
or completely fused into a flat band. The pleural and pedal ganglia are always 
separate, but may lie close together. The visceral pair (supra-intestinal or parietal 



REVISION OF THE WORLD SPECIES OF APLYSIA 285 

and infra-intestinal plus visceral ganglia) are either distinct or fused, but may 
vary with age. They are, however, always on long pleuro-visceral cords and lie 
close to the large hermaphrodite duct. 

Killing and Fixation 

Aplysia is a strongly contractile animal and may shrink to as little as one-third 
of its length when preserved. Much of this shrinkage may be avoided by narcotiza- 
tion before fixing. Three methods are given. 

1. Sprinkle menthol crystals on the surface of the sea water in which the animals 
are relaxed in a vessel of convenient size. When immobile, after about 12 hours, 
fix in 2% chromic acid for 24 hours. Wash in water for 24 hours. Preserve in 5% 
formalin or formol alcohol. 

2. Macginitie's method. Dissolve 75 grams of magnesium chloride and 10 grams 
of urethane in one litre of distilled water. Place the animal in a vessel with sea 
water and add the above solution by the drip string method. When immobile 
proceed as in 1. 

3. If chromic acid and the other ingredients are not available, fix in formol alcohol 
or formalin. 

Formalin hardens the tissues, and in time dissolves the calcareous lining of the 
shell, but it preserves much of the pigment. Alcohol does not harden so much, 
but extracts the colour rapidly. 

Dissection 

It is most important that examination of internal structures should be done 
without disturbing the external appearance of the specimen any more than is neces- 
sary. The internal parts it is necessary to examine are the shell, penis and penis 
sheath, jaws, radula, opaline gland, visceral ganglia, nerve ring and caecum. 

Shell. The mantle aperture should be preserved. Make a cut near the periphery 
of the mantle anteriorly and continue in a crescent along the left side. When the 
cut has extended as far as the shell apex, the shell can usually be lifted out gently 
by raising the flap of mantle tissue. After inspection the shell can be replaced. 

Internal Structures. Examine from the ventral side. Make a longitudinal cut in 
the middle line of the foot and continue forwards through the anterior pedal groove 
to the mouth aperture. Press the cut edges of the foot apart and find the opaline 
gland on the left (animal's right) immediately to the left of the genital duct. To the 
right of the genital duct lies the large colourless cephalic artery and between the 
latter and the globular Swammerdam's vesicle are the supra-intestinal (parietal) 
and visceral ganglia, side by side, and linked by the pleuro visceral cords with the 
nerve ring around the oesophagus anteriorly. To expose them the digestive gland 
and gizzard may have to be turned to the right. 

To find the caecum, turn the animal with the anterior end nearest to you (Text- 
fig. 8b). Remove the covering of connective tissue from the visceral mass, and note 
the positions of the gizzard (right), hermaphrodite gland (posterior) and the course 



286 REVISION OF THE WORLD SPECIES OF APLYSIA 

of the intestine (left). Within this triangle the caecum comes to the surface, either 
lying flat on the digestive gland, or exposing its tip. It may be traced back to the 
stomach by separating the digestive gland tissue. 

Nerve ring (Text-fig. Sa). This consists of cerebral, pleural and pedal ganglia on 
each side, and from the cerebral ganglia connectives run to a pair of buccal or 
stomato-gastric ganglia on the ventral side of the oesophagus, between it and the 
buccal mass. Isolate these ganglia from the buccal mass and oesophagus by cutting 
the nerves, leaving the ganglia attached to the ring by their cerebro-buccal connec- 
tives. Sever the oesophagus close to the buccal mass, and slip it through the nerve 
ring, which can then be examined. Push the buccal mass forward, severing it from 
the body around the mouth aperture. 

Jaws and radula. Boil the buccal mass in a porcelain dish in 5% Potassium 
hydroxide until the jaws and radula can be removed from the softened muscle. A 
small piece cut off the edge of the jaw is sufficient for the examination of the jaw 
rods. The radula should be brushed gently to remove the radular membrane from 
the posterior rows of teeth. After examination under the microscope the separated 
parts can be placed in a small tube to be put into the jar containing the specimen. 

Penis and penis sheath. These lie alongside the buccal mass, on its left side when 
viewed ventrally. The sheath should be slit open longitudinally, so that the penis 
protrudes ; it can then be examined in situ. 

After examination the specimen, closed up again along the foot incision, resumes 
its original shape. 

(e) Criteria Employed in the Description 

Owing to the vagueness of such terms as large, broad, etc. it has been found 
necessary to adopt limits of size, according to the following criteria : — 

Body size : Large — More than 150 mm. preserved, alive 300 mm. or more. 
Medium — 75 to 150 mm. long. 
Small — Up to 75 mm. long. 

Foot : Broad — More than 25% of the body length. 

Medium — 15% to 25% of the body length. 
Narrow — Below 15% of the body length. 

Shell : Broad — Length less than ij times the breadth. 

Narrow — Length at least ij times the breadth. 

It should be noted that the above limits are a guide only and are not absolute. 

In the account of the species which follows, a full list of synonyms, as far as is 
ascertainable, with references, comes after each specific heading. At the end of the 
description the specific characters necessary for identification are given. The list 
of references to the species excludes those given in the synonymy. Full titles of 
papers appear at the end. 



REVISION OF THE WORLD SPECIES OF APLYSIA 287 

2. DESCRIPTIONS OF THE SPECIES 

Subgenus Pruvotaplysia Engel, 1936 

Aplysia (Pruvotaplysia) parvula Guilding in Morch, 1863 : 22 

Text-figs. 10 and 11 

Synonymy : allochroa Bergh, 1908 : 4 (non O'Donoghue, 1929). 

anguilla Cuming, MS. Sowerby, 1869 : pi. 6, shell only. 

atromarginata Bergh, 1905 : 8. 

australiana Clessin, 1899 : 22, shell only. 

concava Sowerby, 1869 : pi. 6, shell only. 

intermedia Farran, 1905 : 351. 

japonica Sowerby, 1869 : pi. 5, shell only. 

lobata Bergh, 1908 : 8 (or this may be a young dactylomela) . 

monochroa Bergh, 1908 : 13. 

nigrocincta Martens, 1880 : 307. 

norfolkensis Sowerby, 1869 : pi. 10, shell only. 

orient 'alis Sowerby, 1869 : pi. 5, shell only. 

paeteliana Clessin, 1899 : 96, shell only. 

peasei Pilsbry, 1895 : 95 as Tethys. 

? peregrina Thiele, 1925 : 281. 

rosea; Sowerby, 1869 : pi. 6, shell only {non Rathke, 1799). 

sieboldiana Clessin, 1899 : 54, shell only. 

subquadrata ; Sowerby, 1869 : pi. 9, shell only (non Gould). 

? trigona Sowerby, 1869 : pi. 4, shell only. 

Aplysiopsis juanina Bergh, 1898 : 483. 

? Syphonota elongata Pease, i860 : 24. 

? Syphonota punctata Pease, 1868 : yy. 

It is possible that Meinertzhagen's tryonii (1880 : 270) belongs to this species, but 
the description is inadequate and the type has been lost. A specimen in Wellington 
Museum labelled tryoni proved to be parvula. 

Krauss in 1848 gave an imperfect description but no figure of a " spurious Aplysia " 
from South Africa, to which he gave the name Aplysia spuria. He compared it 
with Rang's maculata and thought it had no posterior tentacles, hence the name 
spuria. Macnae (1955 : 235) claimed that this is the species later called parvula, 
and that the name should be changed to spuria. There is considerable doubt 
regarding Krauss's species, and the type has disappeared from the Swedish National 
Museum in Stockholm. Application was accordingly made to the International 
Commission on Zoological Nomenclature (April 1957) to validate the specific name 
parvula Morch, 1863, and suppress spuria. This was confirmed in Opinion 560, 
1959- 



288 REVISION OF THE WORLD SPECIES OF APLYSIA 

Pease's species should be ignored, but Ostergaard (1955) has used the name 
Tethys elongata for a Hawaiian species, which may or may not be parvula. Unfor- 
tunately I have been unable to obtain a specimen. 

Distribution. The species is circumtropical in distribution, occurring in warm 
seas from approximately 40 north latitude to 40 south latitude. It has been 
recorded in the western North Atlantic from Florida to Brazil and from many 
islands in the West Indies ; in the eastern North Atlantic from Morocco, the Azores, 
Madeira and Senegal ; in the Indian Ocean from the Gulf of Aden, India, Ceylon 
and Mauritius to the Cape ; in the Pacific Ocean from Japan, China, the East Indies 
to Samoa, the Hawaiian Islands, the Society Islands, Australia and New Zealand ; 
and in the eastern Pacific from California, South California and the west coast of 
South America. 

Material examined. Numerous specimens from all parts of the world. 

This beautiful but variable little Aplysia was described by Morch from two speci- 
mens collected on the island of St. Thomas in the Lesser Antilles, West Indies. 
He stated that he had seen a shell of the species in the British Museum (Natural 
History) from St. Vincent, named by Guilding A. parvula, but no trace of that 
portion of Guilding's manuscript referring to this species can be found. Morch 
noted its resemblance to A. longicornis Rang (— punctata Cuvier, 1803). A. 
parvula was later found in many other parts of the world and appears under local 
names, such as norfolkensis. Engel (1927 and 1936) was the first to show that the 
species is circumtropical. In 1931, Risbec, describing a specimen from Morocco, 
gave a figure of the shell of the young animal with its spire, sketched the characteristic 
wavy contraction of the edges of the foot and the distinct, rounded ganglia of the 
nerve ring. Mme. Pruvot-Fol recognized the primitive characters of the species, 
and in her honour Engel (1936) suggested the subgeneric name Pruvotapiysia. 
Baba (1949, pi. 3), published an excellent coloured reproduction of the living animal 
belonging to the variety nigrocincta. In 1956 he described the egg strings as pink 
or yellow, with only two to three eggs in each capsule. 

Description of the Species 

Small or very small Aplysias, rarely reaching more than 60 mm. in length, although 
the tail may elongate still further. Body rounded and neat, typically higher than 
wide when contracted, but elongated and flexible when moving (Text-figs. 10 and 
11a). Skin soft. Colour very variable — purplish black, dark brown, tan, yellow, 
greenish grey or green, speckled with white or cream. The variety nigrocincta has 
the mantle foramen, siphon, parapodia, rhinophores and cephalic tentacles, anterior 
border of the foot and tip of the tail bordered with black and orange. Another 
variety from Madagascar, now in the Swedish National Museum, has scattered 
papillae on the sides of the body, similar to those figured by Quoy & Gaimard in 
their A. tigrina (= tigrinella M. E, Gray, 1850, = maculata Rang, 1828, according to 
Macnae, 1955). 



REVISION OF THE WORLD SPECIES OF APLYSIA 289 

Head small and neat, neck long. Cephalic tentacles strongly contractile, but when 
expanded large, dilated, sometimes fimbriated, usually free from the mouth slit. 
Eyes plain, usually on light patches. Rhinophores small, slender, acute, split for 
two-thirds of their length, fairly close together. Both the cephalic tentacles and the 
rhinophores are black in the folded portions. 

Foot narrow, pale, with slender, pointed tail ; the edges when contracted have 
a wavy border. Penis (Text-fig. 11c) broad at the base, short and tapering, blunt- 




Fig. 10. Aplysia parvula. Lateral, ventral and dorsal views of living specimens, from 
colour photographs taken by Dr. H. Lemche at Cronulla, New South Wales. The heavy- 
lines indicate the black edges of the cephalic tentacles, rhinophores, anterior and 
posterior borders of the foot, mantle aperture, anal siphon and parapodia. x ij. 

tipped, sheath smooth-lined. Parapodia short, not very mobile, rounded anteriorly, 
widely open, fused posteriorly, forming a moderately high wall round the mantle 
cavity. MacFarland (1924) described them as large and fleshy in Calif ornian speci- 
mens. 

Mantle thin, with a large oval foramen, at least one-third the width of the shell. 
Anal siphon of moderate size. Purple glands well developed. 

Mantle cavity closed behind. Ctenidium small and compact. Opaline gland 
simple, multiporous, though some ductules may coalesce anteriorly. Genital aperture 
rather far forward, covered by the mantle overhang, smooth, crescentic. 

Shell (Text-fig. 116) large for the size of the animal, oval or almost circular, strongly 



290 



REVISION OF THE WORLD SPECIES OF APLYSIA 



concave, thin and delicate in the horny portion, but well calcified beneath, especially 
near the apex, which is hard, incurved, often with a distinct spire especially in young 
specimens, and a small projecting, thickened hood. This is the only species of 
Aplysia which may retain the spiral apex in the adult, although it is not unusual 
for juveniles of other species to exhibit this feature (cf. Marcus, 1955 : pi. 4, fig. 29, 
for Juliana) . Solution of the calcareous portion of the shell, however, reduces the 
apex considerably. 

Jaws triangular. Rods densely packed, slightly curved or straight. 



mant. ap. 



an.s 




b) 



an. si. 




apex 





d) 



+ 1 



+6 



pn.s 




ret.pn 



Fig. 1 1 . Aplysia parvula. 



(a) Right lateral view of a contracted specimen from the Swedish National Museum, 
Stockholm. The specimen shows the high humped appearance of the contracted 
animal, but is unusual in that the sides of the body bear scattered papillae, which are 
more crowded on the tail to form a small keel, x i\. 

(b) Shell in ventral and lateral views, and enlarged in ventral view, from a young specimen 
figured by Risbec, 1931, to show the spiral apex. The dotted line represents the 
limit of the calcareous lining. 

(c) Penis sheath opened longitudinally to show the short broad penis and sperm groove. 
X 6J. 

(d) Radular teeth. Rhachidian, first and sixth laterals, x 80. 



Radula (Text-fig. nd) with few rows and a small number of teeth in a row, a 
typical formula being 30 X 16.1.16. Rhachidian tooth with unusually short but 
wide, incised basal plate, finely denticulate main cusp and one or two stronger 
denticles laterally. First lateral with rather long cusp, similarly denticulate, the 
large lateral denticles increasing in size further along the row, but the cusp remaining 
shorter than the plate. From three to four outermost teeth degenerate. 

Caecum short, just breaking surface on the digestive gland. The hermaphrodite 
gland is pushed up tightly into the spire of the shell. 



REVISION OF THE WORLD SPECIES OF APLYSIA 291 

Nervous system primitive in that all the ganglia of the nerve ring are distinct, 
the pleuro visceral cords are short and the visceral ganglia separate. 

Specific Characters 

Aplysias of small size, neat appearance. Elongated in shape, but of high aspect 
when contracted. Foot narrow, with wavy borders when contracted, parapodia 
joined high up posteriorly. Penis short, with blunt tip, sheath lining smooth. Mantle 
with large oval foramen. Purple secretion from the mantle glands. Opaline gland 
simple, multiporous. Shell relatively large, deeply concave, with strongly inflected 
apex or with distinct spire. Radula with not more than 20 lateral teeth, rhachidian 
tooth with wide but very short basal plate. All the nerve ganglia distinct, pleuro- 
visceral cords short. Colour variable, but an edging of black and orange pigment 
is characteristic of the variety nigrocincta. 

Circumglobal in warm seas. 

References {see also synonyms) 

Allan, Joyce, 1932 : 424, as norfolkensis . 

Angas, 1877 : 190, as norfolkensis. 

Baba, 1937 : 2 °8- 

Baba, Hamatani & Hisai, 1956 : 216 (breeding habits). 

Burne, 1906 : 56, 57, as nigrocincta and elongata. 

Cotton & Godfrey, 1933 : 96, as norfolkensis. 

Dall, 1885 : 24. 

Dall & Simpson, 1902 : 366. 

Edmonson, 1933 : 154, as ? Tethys elongata. 

Eliot, 1899 : 513, as nigrocincta. 

Engel, 1927 : 90 ; 1936a : 15. 

Haas, 1920 : 56. 

Macnae, 1955 : 235, as spuria. 

MacFarland, 1924 : 398, as Tethys. 

Ostergaard, 1955 : 114, as elongata ?. 

Pilsbry, 1895, as parvula, anguilla, elongata, peasei, japonica, nigrocincta, norfolkensis , concava, 

orientalis and trigona, all under Tethys : 83, 112, 93, 95, 106, 107, 99, 100, 104, 112. 
Pruvot-Fol, 1932 : 4 ; 1933 : 104 ; 1935 : 254 (suppression of Bergh's Aplysiopsis). 
Risbec, 193 1 : 74. 

Suter, 191 3 : 545 and pi. 36, fig. 6, as T. tryoni, which is probably parvula. 
Thiele, 1910 : 123. 



Aplysia (Pruvotaplysia) punctata Cuvier, 1803 : 295 

(Fully illustrated by Eales, 1921, q.v.) 

Synonymy : albo-punctata Deshayes, 1853 : 59. 

cuvieri and cuveriana delle Chiaje, 1822 : 41. 
Udepilans ; Pennant, 18 12 : 42. 
depilans minor Barbut, 1783 : 32, pi. 3, fig. 5. 
dumortieri Cantraine, 1840 : 71. 



292 REVISION OF THE WORLD SPECIES OF APLYSIA 

ferussaci Rang, 1828 : 66, pi. 19. 

guttata Sars, M., 1840 : 213, Taf. 7. 

hybrida Sowerby, J., 1806 : pi. 53. 

longicornis Rang, 1828 : 66. 

marginata de Blainville, 1823 : 326 ; Rang, 1828 : 59 ; Philippi, 

1844 : 98. 
mustelina? Davies in Pennant, 1812 : 29, pi. 22. 
nexa Thompson, 1845 : 313. 
nigromarginata Risso, 1818 : 374. 
^rosea Rathke, 1799 : 85 ; Odhner, 1907 : 16 and 61 ; 1922 : 6 

(non Sowerby, 1869) (juv.). 
stellata Risso, 1818 : 374. 

subquadrata Gould in Sowerby, J., 1806 : 111, pi. 53. 
unicolor Risso, 1818 : 374. 
varians Leach, 1847 : 2 ^8 and 1852 : 33. 
virescens Risso, 1826 : 42. 
2 ? Esmia griffithsiana Leach, 1847 : 268 (juvenile) (spelt griffithsiae 

by M. E. Gray, 1850). 

Distribution. Atlantic coasts from Greenland, Norway and the Baltic to the 
Canary Islands and the Mediterranean. The common British species. 

Material examined. Numerous specimens from British and French coasts 
and the Mediterranean. 

A good figure was given by Rang (1828, pi. 18). A full account of the anatomy 
was published by the author (1921). 

Aplysias of moderate size, reaching 200 mm. when alive, but usually smaller. 
Body capable of great elongation, especially in the head and neck regions, contracting 
to the " sitting hare " position. Colour when mature a rich velvety olive brown 
to olive green, resembling the weed Laminaria and Fucus respectively, often accom- 
panied by black spots and reticulations, irregular greyish flecks or white blotches 
formed by the fusion of clusters of unpigmented spots. When preserved the soluble 
brown colour is extracted, leaving black, grey and white. Young specimens about 
6-10 mm. long are rose red, the animals feeding at this stage on red seaweeds such 
as Delesseria. This was described by Garstang in 1890. Skin soft but firm. 

Head and neck long and narrow. Cephalic tentacles short with rounded extensions 
to the mouth slit. Eyes plain, on clear, circular areas of skin. Rhinophores elongated, 
tapering, slit half-way. 

1 Odhner (1907) pointed out that rosea should have priority over Cuvier's punctata. The International 
Commission on Zoological Nomenclature, Opinion 355, 11, 91-102, Aug. 1955, " suppressed the name 
rosea Rathke, 1799, and placed the specific name punctata Cuvier, 1803, as published in the combination 
Aplysia punctata on the Official List of Specific Names in Zoology." 

2 It is not clear whether Leach's specimen was a juvenile punctata or depilans. Although Leach's 
Mollusc collection was given to the B.M. (N.H.) and the specimens were re-named by J. E. Gray, his 
Esmia cannot be found, 



REVISION OF THE WORLD SPECIES OF APLYSIA 293 

Foot narrow, colourless, with rounded anterior edges and long pointed tail, 
adapted for crawling up the stipes of Laminaria, the lateral borders curling towards 
each other. Penis spatulate, flat, sheath smooth-lined, with a single retractor muscle. 
Parapodia mobile, thin, natatory, smooth-edged but capable of forming a fluted 
pattern when the animal is feeding on a frilled weed such as Chondrus crispus, 
joined high up posteriorly, thus closing the mantle cavity behind. 

Mantle thin with a flat circular or oval aperture, tall, tubular anal siphon and well 
developed purple gland. 

Mantle cavity not large, opaline gland simple, multiporous, but small groups of 
vesicles may fuse. Genital aperture smooth, pigmented. 

Shell delicate, transparent, moderately concave, rounded anteriorly, apex down- 
turned but not spiral, anal sinus very shallow, dorsal border of apex reflected. 
In the young, the spire of the shell, as in many juvenile Aplysias, is well formed, 
cf. Mazzarelli, 1893 : pi. 12. 

Jaw rods of moderate length, straight, but tapering and slightly bent at the tip. 

Radula small, not heavily chitinized. About 40 rows and a formula 15-18 . 1 . 15-18. 
Rhachidian tooth with broad, fairly short basal plate, incised both anteriorly and 
posteriorly, cusp narrow near the point, a little longer than the plate, bearing a 
few small neat denticles and two pairs of large basal ones, the penultimate denticle 
being the larger of the two. First lateral with narrowing, almost triangular basal 
plate, broad head, narrow cusp repeating the rhachidian pattern laterally. The larger 
denticle may split into two or more. Along the row the plates remain narrow, the 
heads and cusps become more slender, but the cusps do not exceed the length of 
the plates, the denticles increase in size, particularly the lateral basal ones. The 
two outermost teeth are small and vestigial. 

Caecum short, only the curved tip visible on the surface of the digestive gland. 

Cerebral, pleural and pedal ganglia rounded and distinct. Visceral group (right 
parietal and visceral) also separate. 

Specific Characters 

Of moderate size, with rich velvety olive brown or olive green colour, with or 
without black reticulations and clear areas, and soft firm skin. Head and neck long, 
but contracting back to the visceral hump to give a " sitting hare " appearance. 
Foot narrow, unpigmented, with pointed tail. Penis broad, spatulate, flat. Parapodia 
natatory, joined high up posteriorly. Mantle with flat aperture. Purple secretion. 
Opaline gland simple, multiporous. Shell delicate, with weak anal sinus, inturned 
apex, but no spire. Radula small, with neat regular denticles and a formula 40 x 
18. 1. 18. All the ganglia of the nerve ring rounded and separate, as are also the 
visceral ganglia. 

Eastern North Atlantic. 

References (see also synonyms) 
Colgan, 191 1 : 21. 
Dautzenberg & Fischer, 1925 : 29. 
Eales, 1921 : 1-84. 
Engel, 1934 : 85-89. 



294 REVISION OF THE WORLD SPECIES OF APLYSIA 

F6russac, 1821-22 : xxx. 

Fischer-Piette, 1932 : 14. 

Fleming, 1828 : 291. 

Garstang, 1890 : 401. 

Grigg, 1949 : 795- 

Guerin-Meneville, 1843 : 20. 

Hertling, 1935 : 103. 

Hoffman, 1926 : 7. 

Jeffreys, Gwyn, 1866 : 81 ; 1869 : 1. 

Locard, 1886 : 59. 

Mazzarelli, 1889 : 582 ; 1893 : 25. 

Norman, 1890 : 69. 

Pilsbry, 1895 : 70, as Tethys. 

Saint Loup, 1888 : 1010. 

Sars, G. O., 1878 : 213. 

Sars, M., 1878 : Tab. 12, fig. 18. 

Sykes, 1905 : 78. 

Vayssiere, 1885 : 54 ; 1935 : No. 20 (not paginated). 

Verany, 1853 : 390. 

Note. Some specimens may be hybrids. Gwyn Jeffreys (1866) saw Aplysia 
depilans and A. punctata copulating in a tank in the Channel Islands. This may 
explain some of the intermediate forms, e.g. A. lobiancoi Mazzarelli, 1890 : 42. 



Subgenus Neaplysia Cooper, 1863 

Aplysia (Neaplysia) calif ornica Cooper, 1863 : 57 

Text-figs. 12 and 13 

Synonymy : A. ritteri Cockerell, 1901-02 : 90. 
? A. nettiae Winkler, 1959 : 8. 1 

Distribution. West coast of North America : California and Lower California, 
including the Gulf of California. 

Material examined. Several specimens from California (B.M. (N.H.)) and the 
Gulf of California, the latter sent by Dr. L. R. Winkler. 

Pilsbry (1895, pi. 56) gave a figure of the whole animal. The external features 
and the shell were described by Cooper, who founded the subgenus on the peculiar 
flattening of the shell apex, which he regarded, probably incorrectly, as an " acces- 
sory plate ". 

One of the giant members of the genus, attaining 375 mm. in length, 125 in breadth 
and 125 in height. Macginitie (1934) recorded a specimen weighing 5 lb. 12 oz. 

1 Winkler's preserved specimens, described as a new species, are probably a variety. His account is 
not detailed enough to justify founding a new species on such slender grounds as open mantle tube, 
more frilly parapodia, and dark colouration, although the separation of the posterior attachments of the 
parapodia is a distinctive feature. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



295 



which laid 478 million eggs in 18 weeks. It is a plain species, with rather low, 
swollen body, thick in front, with the mantle and parapodia far back, the foot 
terminating in a long slender tail. It resembles punctata when it assumes the " sitting 
hare " position. The skin is soft, smooth and flabby. Colouring is very variable. 
Pale grey, green or dark brown, marbled with crimson or purple on the sides, and 
with a network of fine brown or black lines all over, usually with numerous large 
brown blotches and dark brown or black irregularly placed spots. Where these 
round spots lie on the intersections of the reticulations they may produce the illusion 



o) 



b) 







Fig. 
specimen in 



12. Aplysia calif ornica. 

dorsal and lateral views, x f . Calcareous area 



{a, b) Shell of one 
dotted. 

(c) Shell of a larger specimen, in which the calcareous and horny layers are distinct. It 
is evident that the " accessory plate " of Cooper (1863) is the flattened apical portion 
of the calcareous layer, which in (a) and (b) projects as a rectangular plate, x 2. 

(d) Penis sheath in ventral view, with its two retractor muscles, x 1. 
{e) Penis, x 2. 



of a star pattern. The inner sides of the parapodial edges have a greenish back- 
ground and bars of brownish black alternating with lighter areas, a common pattern 
in the genus. Mantle clear or with white flecks. Foot dark to black. All colour and 
markings may disappear with preservation. 

Head small but broad and clumsy, strongly contractile. Cephalic tentacles short, 
folded, lined with black and linked with the ventral sides of the mouth slit. Rhino- 
phores close together, stout, black within the short slit ; when contracted they 
appear cup-shaped. Eyes small. 

Foot moderate to broad, carunculate, rounded anteriorly, with defined wrinkled 
edges and a long slender tail. The animal attaches itself by the anterior and posterior 
ends of the foot, as many of its relatives do, raising the foot sole between these 



296 REVISION OF THE WORLD SPECIES OF APLYSIA 

two points, but a sucker is not defined. Penis broad, tapering at the apex, coiled 
in the bulbous smooth-lined portion of the sheath (Text-fig. 12, d, e). Parapodia 
short, but can be raised high ; they arise close together, have slightly wavy, thin, 
freely mobile edges, and are joined posteriorly low down, forming a small platform 
over the foot. 

Mantle small with a minute, usually closed tube, sometimes marked with concentric 
wrinkles. Anal siphon broad, thin, tubular, frilled on the edges and capable of 
considerable extension. A deep purple secretion from the mantle glands. 

Mantle cavity widely open at both ends. Ctenidium small, unpigmented. Opaline 
gland large, compound, with one aperture, sometimes raised on a papilla. Genital 
aperture small, smooth, crescentic. 





Fig. 13. Aplysia calif ornica. Radular teeth. Rhachidian and first, sixth, 
and outermost laterals, x 80. 



The shell (Text-fig. 12, a-c) is broad and short, rounded in shape, but narrowing 
almost to a point near the anterior edge. It is only slightly concave, without a 
spire and with very little calcareous matter. The growth lines are plainly visible 
and there are also radial striae. The anal sinus is weak. There is a recurved dorsal 
edge at the apex, and on the ventral side a rectangular plate, which Cooper described 
as an " accessory plate ". Examination of several recently preserved individuals, 
however, shows that this plate is the flattened calcareous portion of the apex, which 
separates from the horny apex. It is, therefore, a part of the shell, and not an 
accessory structure. 1 

The jaws consist of tall slender curved rods. 

The radula in two small specimens had 50 rows and 56 rows and a formula of 
32 . 1 . 32 and 40 . 1 . 40 respectively, but large specimens must exceed this considerably 
(Text-fig. 13). The rhachidian tooth has a broad basal plate, straight posteriorly, 
rounded at the corners, slightly waisted and with a narrow and deeply incised head. 
The cusp is shorter than the plate in mature teeth, a little longer in young teeth, 
with small irregular denticles, the basal one being short and divided. The first 
lateral has a rounded out- turned plate, broad rounded head and similar irregular 
denticulations. The slender cusp of the thirteenth tooth reaches the maximum 
length in the row, and the denticulations increase correspondingly, while the bases 
become more slender. There is a tendency for large denticles to become subdivided. 
The six outermost teeth are without cusps. 

1 According to Winkler (1958), who examined young individals, the accessory plate "contains the 
built-in veliger and the metamorphic shell." 



REVISION OF THE WORLD SPECIES OF APLYSIA 297 

The caecum is very long, with enlarged recurved tip. 

The cerebral ganglia form a broad flat band. The visceral ganglia are joined. 

Specific Characters 

Large Aplysias, with soft skin, irregularly mottled with brown and with fine 
reticulations. Shape clumsy, visceral region small, foot broad with long slender 
tail, parapodia short, mobile, joined low down posteriorly, mantle closed, with 
papilla, purple glands, broad flat shell with rectangular flattened calcareous apex 
and no spire, radula with elaborate but irregular dent iculat ions, penis broad, taper- 
ing, opaline gland compound, uniporous. 

California. 

References {see also synonyms) 
Cockerell, 191 5 : 84. 
Cooper, 1867 : 14. 
Lowe, 1934 : 43. 
Macginitie, 1934 : 3°°- 
Parker, 191 8 : 139. 
Pilsbry, 1895 : 89, pi. 56, as Tethys. 
Winkler 1958 : 348 ; 1959 : 8. 

Subgenus Varria no v. 

Aplysia (Varria) brasiliana Rang, 1828 : 55, pi. 8 

Text-figs. 14 and 15 

Synonymy : Tethys floridensis Pilsbry, 1895 : 82. I have seen the type, and 
could find no significant difference in structure between it and 
Rang's species. 

A. livida d'Orbigny, 1837 : 206. Mis-spelt lurida on plate 18. 
A. guadeloupensis Sowerby, 1869 : pi. 5, named from the shell 

only, may be this species. 
A. cailleti Deshayes, 1857 : 140, probably this species. 

Distribution. The species belongs to the Atlantic Ocean, and ranges from New 
Jersey in the north to St. Helena in the south. It has been found in Florida, Texas, 
the West Indies and Brazil, and on the coast of Ghana in West Africa. 

Material examined. Numerous examples from the eastern seaboard of North 
and South America and from West Africa. 

Rang figured brasiliana and his figure was copied by Pilsbry. Marcus (1955) has 
published an excellent account of it and has studied the development. He can 
find no valid distinction between brasiliana and d'Orbigny's livida. I am indebted 
to Dr. Marcus for observations on the living animal and for co-operation in preparing 
the following account. 

zooi.. 5, 10. 15 



298 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Medium to large Aplysias, high and massive, with elongated head and neck, 
but strongly contractile. Much swollen posteriorly. Reaching 270 mm. when alive, 
and from 140 to 170 mm. high. Skin smooth, but rugose when preserved, " like 
peccary leather ". Colour usually yellow, deep brown, bottle green, grey or purplish 
black, unicoloured or spotted yellow, dark brown or green, with black pigment 
forming a fine veining or network. There may be white or dark spots or chalky 
granules, connected by a network. Occasionally creamy white specimens occur. 
A very variable species. The parapodia are blotched with black on their inner 
surfaces, with alternately dark and light areas arranged vertically near the edges, 
which are light. Foot dark brown to light brown, usually uniformly coloured. 




Fig. 14. Aplysia brasiliana. Shell of two specimens in ventral 
and dorsal views respectively, x f . 



+12 




Fig. 15. Aplysia brasiliana. Radular teeth. Rhachidian and first, sixth, 
12th and outermost lateral teeth, x 80. 



Mantle dark with radiate or irregular light areas. Ctenidium usually pigmented. 
Tips of the cephalic tentacles and rhinophores light. 

Head and neck elongated, small. Cephalic tentacles broad, but not meeting mid- 
dorsally. Eyes in clear patches. Rhinophores small, conical, set close together, 
slit half-way down. 

Foot narrow to medium, with a short tail. Penis long, broader at the base, tapering 
but not filiform, penis sheath smooth. Parapodia large, can be widely opened 
exposing the whole of the dorsal area, joined posteriorly very far back and low down 
on the root of the tail, the edges upstanding before joining. (Rang's figure is mis- 
leading.) The edges are thin and wavy, but not fimbriated. 

Mantle large, set far back, thin. Mantle aperture minute, on the tip of a papilla. 
Anal siphon of moderate to large size, plain or wavy, sometimes projecting backwards, 
wide but not high. Mantle glands secrete a magenta red secretion. 

Mantle cavity open at both ends. Ctenidium large, but contractile. Opaline 
gland compound, uniporous. Marcus says that the secretion is greenish yellow but 



REVISION OF THE WORLD SPECIES OF APLYSIA 299 

not fetid. Genital aperture smooth or furrowed, pigmented, set well under cover 
of the mantle. Genital groove plain. 

Shell (Text-fig. 14) narrow to broad in large animals, concave, dark yellow, with 
the apex hard and hooked, but without spire. Anal sinus shallow, firm edged, 
dorsal part of apex recurved. Usually with marked concentric rings and one or 
more radial striae. 

Jaws very dark. Rods long, slender, almost straight, but some may have bent 
tips with fringed edges. 

Radula almost as broad as long, 65-70 rows, formula 43.1.43 (Text-fig. 15). 
Rhachidian tooth narrow, with long plate and narrow head, simple cusp bearing 
one basal denticle on each side and a few irregular ones. First lateral small, with 
rectangular plate and similar denticulations. Remaining teeth with longer, narrower 
cusps, a pair of lateral denticles, and the other denticles so small that the edge 
appears beaded, except in young teeth. Further along the row the plates become 
attenuated and the cusps longer. About three terminal teeth are degenerate. 

Caecum appears on the surface and is straight and inflated at the tip. 

Cerebral ganglia band-like, fused. Visceral ganglia joined. 

Specific Characters 

Large bulky body, self-coloured or spotted. Simple cephalic tentacles, slender 
rhinophores set close together, long neck, narrow foot, long tapering penis, very 
large parapodia joined low down posteriorly, mantle aperture tubular, opaline 
gland compound, uniporous, simple radula with high narrow rhachidian tooth. 
Young specimens may have a hole in the mantle. 

West Atlantic from New Jersey to Brazil ; East Atlantic : Ghana and Saint 
Helena. 

References (see also synonyms) 

Adams & Adams, 1858 : 35 (as Syphonota lurida). 
Marcus, 1955 : 4. 

Pilsbry, 1895 : 82 and 79 (livida), as Tethys. 
Pruvot-Fol, 1934 : 3^ • 

Aplysia (Varria) cervina Dall & Simpson, 1901 : 365, pi. 56, fig. 2 

Text-figs. 16 and 17 
Synonymy : pilsbryi Letson, 1889 : 193. Yucatan. 

Distribution. Atlantic seaboard from Eastern U.S.A., through Central America 
and the West Indies to Brazil. 

Material examined. Numerous specimens from eastern U.S.A. and Central 
America, including two ' A. pilsbryi,' belonging to the Academy of Natural Sciences, 
Philadelphia. 1 

1 Except for an upturned edge to the rim of the mantle where the purple gland lies, to which Letson 
referred, there is no structural difference between these specimens and cervina. 



3°° 



REVISION OF THE WORLD SPECIES OF APLYSIA 



The species was named by Dall & Simpson from nine rather small specimens 
from Porto Rico. MacFarland (1909 : 38) gave a long account of its anatomy. 

Of moderate size, smaller, more bulky and plainer than brasiliana, reaching about 
70 mm. when preserved, but more elongated when alive (Text-fig. 16a). Soft, 




b) 



% 






gvf<> 



00 *£8 - . 



c) 








Fig. 16. Aplysia cervina. 



(a) Dorsal view of a specimen from Anguilla, West Indies, x f . 

(b) Pattern on the side of the body, composed of round dark brown spots and clear areas, 
sometimes in clusters or rosettes on a light brown background, which is finely reticu- 
lated, but the reticulations have not been figured. 

(c) Pattern on the inner side of a parapodium, made up of dark and light blotches. 

(d) Penis sheath and penis, x ij. 



plump and smooth, flabby. Grey with scattered round black or smoky brown spots 
and flecks up to 2-5 mm. in diameter, with reticulations and blotches between the 
spots, and there may be lighter areas. A specimen may have about 30 spots on the 
side of the body (Text-fig. 166). Mantle dappled with alternating light and dark 
patches, sometimes arranged radially. Inner sides of the parapodia grey to black, 



REVISION OF THE WORLD SPECIES OF APLYSIA 



301 



with vertical black and light areas, but clear on the edges (Text-fig. 16c). Foot 
dark. All colour and markings may disappear after long preservation. 

Head and neck rather small. Cephalic tentacles plain, short in contracted speci- 
mens, but can be spread like elephant's ears. Rhinophores conical, fairly widely 
spaced, but not as wide as the parapodial origins, deeply slit. Eyes far apart. 

Foot with narrow to medium sole, tough and wrinkled in contraction, the anterior 
border with rounded angles, the tail short, triangular or blunt. Penis (Text-fig. i6d) 
short, broad, flattened, capable of great elongation, may be curled laterally and 
longitudinally, apex acute. Sheath small, with large muscles attached by two strong 
bands to the bulbous portion, the lining of which is smooth. Parapodia rather 
thick, not large, about three-quarters of the length of the animal, the anterior ends 





Fig. 17. Aplysia cervina. Radular teeth. Rhachidian, first, sixth, tenth and outer- 
most laterals. x 80. Below, the cusp of an old rhachidian tooth, enlarged, to show 
irregularity of the denticulation. 



widely separated, edges slightly sinuous, coming close together and joined low down 
posteriorly, forming a flat extension of the mantle floor. 

Mantle thin, with a small aperture on a papilla, sometimes with radiating lines 
around it. Siphon broad, fimbriated, not high. Purple glands present. 

Mantle cavity " roomy " (MacFarland) , opening out behind between the low 
fused parapodia. Ctenidium large. Opaline gland compound with a single aperture 
on a low thickened elevation. Genital aperture oblique, pigmented, the edge some- 
times projecting in a small peak dorsally. Genital groove deep and conspicuous. 

Shell thin, flat, narrow to broad and rounded. Apex recurved dorsally but imper- 
fect ventrally. Anal sinus long and shallow. Calcareous and horny parts present. 

Jaws with rounded edges. 

Radula 38-48 rows, formula 22 . 1 . 22 but an older specimen may have up to 40 
teeth on each side (Text-fig. 17). Rhachidian with broad plate, slightly excavated 
on the posterior border and more deeply so anteriorly, between the two a narrow 
or waisted effect. Cusp longer than the plate, but shortening with wear. Two 
pairs of lateral denticles near the base larger than the others, which are small and 



3 o2 REVISION OF THE WORLD SPECIES OF APLYSIA 

regular, but not deeply cut, becoming irregular with age. First lateral with bulbous 
head, higher on the antero-lateral portion, the remainder of the plate tapering, 
out-turned. In the remaining laterals the cusp does not lengthen greatly, the plate 
remaining bulbous anteriorly and the denticles exhibiting a common type, the 
large basal denticle sometimes splitting or even becoming multidenticulate. The 
last four or five teeth in the row are rounded and weak. 

Caecum curved, lying flat on the surface where exposed. 

Cerebral ganglia completely fused. Visceral ganglia joined but their double 
origin can be seen anteriorly. 

Specific Characters 

Moderate size, plain unfimbriated aspect, with scattered brown spots on the sides 
of the body, vertical bands of dark and light on the inner sides of the parapodia, 
small head and plain cephalic tentacles, short broad flattened penis, narrow foot, 
low joined parapodia, small open papilla on the mantle, compound uniporous 
opaline gland, constricted plate of the rhachidian tooth and bulbous heads of the 
laterals. 

West Atlantic from U.S.A. to Brazil. 

References {see also synonymy) 
Engel, 1936 : 13. 
van Jhering, 19 15 : 135, 140. 
MacFarland, 1909 : 38. 
Marcus, 1959 : 3. 

Note. Van Jhering thought that MacFarland confused brasiliana and cervina 
and that the latter belongs to the West Indies, but his distinction refers to colour 
and markings only. Although there are resemblances the species appear to be 
distinct. 



Aplysia (Varria) cornigera Sowerby, 1869 : pi. 9 

Text-fig. 18, a, b, c 

Distribution : India, Ceylon, Indo-China, the Philippines. 

Material examined. Several specimens from India, Ceylon and the Philippines. 

The species was named from a shell only by Sowerby. Farran (1905) stated that 
it is a common Ceylon species. A description was given by the author (1944 : 2). 

Aplysias of moderate size, usually about 90 mm. long, 66 mm. wide and 75 mm. 
high when preserved, rather bulky. Brownish olive with clear spaces, in the centre 
of which there may be a black spot. Mottled pigment on the inner sides of the 



REVISION OF THE WORLD SPECIES OF APLYSIA 303 

parapodia near the edges. All pigment may disappear with long preservation. Skin 
very smooth and not strongly contractile. 

Head and neck broad and short. Cephalic tentacles flat, leaf-like, wholly lateral 
and slightly rolled at the edges, continued over the mouth. Eyes small, distinct. 
Rhinophores slender, rounded, set well back, close together. 

Foot narrow, defined laterally, with long pointed tail. Penis (Text-fig. 18c) 
short, flat, sickle-shaped, moderately broad at the base, the spermatic groove lying 
along its edge. Parapodia arising about half-way between the rhinophores and the 
genital aperture, united low down posteriorly but above the level of the mantle 
cavity floor, so that there is a shallow mantle cavity pocket. Margins smooth or 
slightly sinuous but not frilled. Dorsal slit wide open. 

Mantle of moderate size. Mantle foramen minute or closed, with radial markings 
around it. Purple glands present. Anal siphon short and broad. 





i] 



Fig. 18. Aplysia comigera. 

(a) Ventral, (b) dorsal view of the shell, x f . 
(c) Penis, x 2. 

Mantle cavity floor almost flat, with a small pocket behind. Genital aperture 
smooth, crescentic, pigmented. Even in small specimens the genital duct shows 
plainly through the floor. Opaline gland simple, multiporous, though there is a 
tendency for several gland cells to join and open together in the anterior part of the 
gland. 

Shell (Text-fig. 18, a, b) thin, delicate, broadly ovate, rounded anteriorly, deeply 
concave. Apex inturned but weak. Anal sinus short and not deep. A specimen 
90 mm. long had a shell measuring 43 X 36 mm. 

Jaw rods short and compact, very dark, curved. 

Radula about 40 rows, formula 30.1.30. The rhachidian tooth has a broad 
triangular basal plate, short cusp with small blunt denticles, the two basal ones 
larger than the others. The laterals have denticulate main cusps, with one larger 
denticle mesially and a group of very irregular denticles laterally, the penultimate 
denticle being the largest, the outermost three to five teeth are vestigial. 

The exposed portion of the caecum lies flat on the surface of the digestive gland. 

The visceral ganglia are partly fused, the cerebral entirely so. 

The species has been compared by Farran (1905) with A. pulmonica Gould from 
Samoa. It is quite distinct, however, and differs from that species in its shape, 



3°4 



REVISION OF THE WORPD SPECIES OF APLYSIA 



narrow foot, multiporous opaline gland, shape of the shell, radular formula and tooth 
pattern. 

Specific Characters 

Moderate size, smooth, mottled and spotted skin, narrow foot and tail, low fusion 
of the parapodia posteriorly with shallow mantle cavity pocket, closed or almost 
closed mantle, simple opaline gland with many apertures, broadly ovate delicate 
shell and denticulate radular teeth. 

Northern part of Indian Ocean and East Indies. 

References 
Eales, 1944 : 2. 
Farran, 1905 : 2. 
Pilsbry, 1895 : 103, as Tethys. 



Aplysia (Varria) cronullae sp. now 

Text-figs. 19 and 20 

Distribution. Australia : New South Wales. 

Material examined. Three specimens from Cabbage Tree Creek, Cronulla, 
collected at low water in 1951 by Dr. H. Lemche, and one specimen, juvenile, from 
Port Hacking, collected in 1914 by Dr. Th. Mortensen. The largest specimen in 
the University Museum, Copenhagen, is designated as the holotype. One of the 
others was presented to the British Museum (Nat. Hist.) by Dr. Lemche. 

Of medium srze, but massive and bulky. Alive, Dr. Lemche states that it measured 
110-140 mm. in length, 40-50 mm. wide and high, the head 25 mm. long and 20 mm. 
broad. The largest preserved specimen measures 106 mm. long, 43 mm. wide and 
58 mm. high, the foot being 27 mm. wide (Text-fig. 19, a, d). Colour greyish green, 
mottled dark brownish green. In addition to the irregular blotches there are fine 
black lines forming a network or running parallel, especially on the head. The inner 
sides of the parapodia have dark blotches, but the edges are light. The mantle 
has radiating stripes of pigment from the central papilla to the edges. Foot mottled. 
The colour is very variable and may be supplemented by white patches. Skin soft. 

Head rather large, neck short and wide. Cephalic tentacles standing out well on 
the sides of the head with the folded part small, then becoming sinuous, with wide 
rounded mouth flaps. Rhinophores round, elongated, pointed, close together, 
slit half-way, the lining of the slit dark. Eyes very small. 

Foot of medium width, soft, the edges defined, the anterior edge rounded, with 
a dark line on the border, tail short and pointed. Penis sheath small, straight, with 
two small retractor muscles. Penis (Text-fig. 19/) narrow, unpigmented, filiform. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



305 



Parapodia well developed, soft, thin-edged but sinuous, rounded, arising about as 
far back from the rhinophores as these are from the cephalic tentacles, meeting 




Fig. 19. Aplysia cronullae. 

(a) Lateral view, showing the blotched skin, but not the lines or reticulations, x § . 

(b) Anterior view of the cephalic tentacles and mouth slit. x 1. 

(c) A rhinophore. 

(d) Dorsal view of the mantle to show the bands of pigment readiating from the central 
papilla. The ctenidium, genital aperture and sperm groove, anal siphon, anus, 
opaline gland apertures, parapodia and tail are figured, x 1. 

(e) Shell in dorsal and ventral views. X f . 

(/) Penis partly protruded, and right, in its sheath, with the two retractor muscles. 



The penis is unpigmented, but has been drawn in black, owing to its small size. 



X I: 



but not joining low down on the tail, forming a platform continuous with the mantle 
cavity floor. 

Mantle (Text-fig. igd) thin with a small open aperture on a minute papilla. Anal 
siphon well developed, broad and short, fluted. Purple glands dark. 

Mantle cavity large, genital aperture pigmented, smooth-edged, the duct lying 



3°6 



REVISION OF THE WORLD SPECIES OF APLYSIA 



straight under the mantle cavity floor. Opaline gland large, more than twice the 
size of that in A. sydneyensis, simple, multiporous, curved, pyriform. 

Shell (Text-fig. 190) 38 x 27 mm., almost flat, with well marked concentric 
ridges, weak apex and slight anal sinus. 

Jaws pale, rectangular, the rods of moderate length, straight or slightly curved. 

Radula (Text-fig. 20) large, with about 50 rows and a formula of 28 . 1 . 28. Rhach- 
idian tooth with wide basal plate, almost straight posteriorly, deeply incised 
anteriorly. Cusp short, rounded, with a few irregular denticles. First lateral with 
triangular basal plate and firm denticulations. Along the row the plates lengthen, 
then broaden considerably, becoming squared off. The denticulations are strong 
but not regular, the cusps of moderate length only. Three teeth are vestigial at 
each end of a row. 



+ 12 




Fig. 20. Aplysia cronullae. Teeth from the radula. Rhachidian, first, sixth, 12th, 
1 8th and four outermost laterals, x 80. A small portion of the blade of a lateral 
near the middle of the row has been enlarged. 



The caecum lies flat on the surface of the digestive gland and may be inflated at 
the tip. 

Nerve ganglia : Cerebral ganglia fused, visceral also. 

Specific Characters 

Medium size, bulky shape, greenish colour with dark brownish green irregular 
blotches, fine black reticulations and parallel lines on the head, the inner sides of 
the parapodia with blotches, but the edges light, mantle with radiating bands of 
dark pigment from the central papilla, sometimes white patches. Skin soft. Cephalic 
tentacles large, rounded, rhinophores slender, set close together. Foot of medium 
width, defined edges and pointed tail. Penis sheath small, penis filiform. Parapodia 
rounded, meeting low down posteriorly. Mantle with a small aperture on a papilla, 
anal siphon broad and short, purple glands. Opaline gland large, simple, multiporous. 
Shell broad, flat, with weak apex and shallow sinus. Radula about 50 x 28.1.28, 
with irregular denticles. Caecum straight. 

Australia, New South Wales. 



REVISION OF THE WORLD SPECIES OF APLYSIA 307 

Aplysia (Varria) dactylomela Rang, 1828 : 56, pi. 12 

Frontispiece and Text-figs. 1-8 

Synonymy : A. d. var. aequorea Heilprin, 1888 : 326. 

angasi Sowerby, 1869 : pi. 8 ; Hedley, 1918 : 107 ; 1923 : 314, pi. 33. 

annulifera Thiele, 1930 : 586. 

argus Riippell & Leuckart, 1828 : 23. 

benedicti Eliot, 1899 : 513. 

fimbriata Adams & Reeve, 1850 : 63. 

lobata? Bergh, 1908 : 148 (? juvenile). See also parvula. 

megaptera Verrill, 1899 : 545 ; 1901 : 26. 

var. ocellata d'Orbigny, 1835-44 : 44> P 1 - 5 ( non Adams, 1861). 

odorata Risbec, 1928 : 51 and 1951 : 123 var. bourailli. 

operta Burne, 1906 : 51. ? Mis-spelt aperta (as Tethys) by Turton 

1932. 
panamensis Pilsbry, 1895 : 88, as Tethys. 
protect, Rang, 1828 : 56. 

radiata Ehrenberg, 1831 (pages not numbered) {non Crouch, 1826). 
schrammii Deshayes, 1857 : I 4°- 
scutellata Ehrenberg, 1831. 
tigrina Rang, 1828 : 57 {non Q. & G.). 
velifer Bergh, 1905 : 12 (juvenile). 
Syphonota viridescens Pease, 1868 : 77. 
PSyphonota keraudreni ; Angas, 1867:227; Sowerby, 1869 {non 

Rang). 
guadeloupensis Sowerby, 1869 : pi. 5 may be dactylomela or 

brasiliana. 

Specimens recently collected and known locally as A. grandis (Pease, i860) were 
sent to me by Dr. Alison Kay, of the University of Honolulu. They proved to be 
dactylomela. The identity of Pease's Syphonota grandis remains in doubt. 

Distribution. Worldwide in warm seas. Recorded from Bermuda, Florida, 
Mexico, the West Indies, Panama, Brazil, the Canaries, Cape Verde Islands, Ghana, 
the Red Sea, Ceylon and India, Mauritius, South Africa, China, Japan, the East 
Indies, the Philippines, Samoa, Tonga, the Gilbert Islands, Australia and New 
Zealand. 

Material examined. Numerous specimens from all areas in its range. 

No species of Aplysia has received more notice than this, owing, no doubt, to its 
large size, wide distribution and striking appearance. Rang, who named it, gave a 
poor figure of it, and recognized protea and tigrina as separate species, although 



3 o8 REVISION OF THE WORLD SPECIES OF APLYSIA 

they are almost certainly the same. The types cannot be found. The species varies 
very little in different parts of the world. It has received local names ; Engel (1929) 
correlated these and showed that dactylomela is circumtropical. A beautiful coloured 
figure, based on a painting by Dr. J. Stuart, was published by Hedley in 1923, and 
has been used for a black and white illustration as frontispiece to this Memoir. 
Baba (1949) also figured it in colour. MacFarland (1909 : 14) gave a lengthy account 
of its structure. 

Large, plump and bulky, attaining more than 400 mm. in length when alive, 
though many specimens do not exceed 300 mm. and the largest contract to this 
length when preserved (Text-fig. 1). Contracts to the sitting hare position, the head 
and neck being drawn back almost to the anterior edges of the parapodia. In exten- 
sion these regions are long and rather flattened. Skin soft, but may become rough 
and tough in contraction. Ground colour dark brown to olive green or yellowish 
green, with light bluish eye flecks surrounded by large, scattered, black or brown 
rings of irregular sizes and badly defined outlines (Text-fig. 2). There are usually 
from 20 to 30 rings on each side of the body, and there may be one on the centre 
of the head. The rings may reach 25 mm. in diameter and 3 mm. in thickness. Some- 
times there are pigment bands between the rhinophores and the cephalic tentacles, 
and there is usually a dark blotch on the dorsal surface of the tail. The rings may 
be prolonged on their outer edges into branched black lines which may become 
reticulate and anastomose with those from adjacent rings. From a study of young 
specimens it is evident that the rings are formed from the fusion of strands of pigment, 
hence the inter-annular connections. The centre of the ring may be traversed by 
similar pigment strands. White opaque dots, probably of a calcareous nature, 
and finer black anastomosing lines may also be present. On the inner sides of the 
parapodia the black pigment is massed in the form of branched vertical bands ; 
the edges are light. There is a tendency for the contracted tentacles, parapodial 
edges and anal siphon to assume a frilled appearance, accentuated by their sinuous 
or fluted margins. The var. fimbriata, common in the Indian Ocean, exhibits this 
character. The foot sole is pale with dark brown irregular blotches. The animal 
is a strong swimmer. 

Head and neck large, but short and squat in contraction. Cephalic tentacles 
large, with smooth or fringed margins, folded as usual, continued to the sides of 
the mouth. Rhinophores thick, set close together, with shallow grooves notched 
on one side, when contracted appear cup-shaped. 

Foot (Text-fig. 3a) broad, thick, soft, but becomes hard in contraction, its edges 
well defined, front edges rounded, tail short and blunt. Penis sheath attached by 
two strong retractor muscles, supplemented by finer strands at the sides (Text-fig. 
5c). Penis white, broad, spatulate, long when extended, the sperm groove winding 
spirally round it. Parapodia (Text-fig. 4) large, powerful, natatory ; they commence 
close together ; edges sinuous or fluted, occasionally smooth, not normally covering 
the anal siphon, joined posteriorly low down on the tail, forming a platform-like 
extension of the mantle cavity floor or at most a low pocket. Sometimes they are 
free, with a pigmented gap between the lobes. 



REVISION OF THE WORLD SPECIES OF APLYSIA 309 

Mantle (Text-fig. 4) large, thin, sometimes rough, with a small foramen, but more 
commonly a closed conical papilla. Anal siphon large and frilled. Deep purple 
mantle gland exudate. A. protea of Martinique, according to d'Orbigny (1853), 
is called locally " Baril de vin " owing to the deep reddish purple secretion it exudes 
when disturbed. 

Mantle cavity shallow, ctenidium of moderate size, usually not pigmented. Opaline 
gland (Text-fig. 5#) compound, forming a large solid trihedral block hanging beneath 
the mantle cavity floor by its duct. Genital aperture smooth, oblique, crescentic 
or with a little triangular peak; marked by a black streak. 

Shell (Text-fig. 6) large, broad, rounded, with an oblique apex, strongly calcified, 
with hardly any trace of a spire, but with a deeply excavated anal sinus. It varies 
with age from thin and delicate to thick and solid. Postero-dorsal edge recurved. 

Jaws (Text-fig. 7, e, f) rectangular, about 5x3 mm., dark in colour. Jaw rods 
short, curved, broadening at their free ends, or straight and truncated. 

Radula (Text-fig. 7, a-d) large. For a specimen measuring 170 mm. Macnae 
gave the formula 73 X 67.1.67. Rhachidian tooth wide, with slightly excavated 
basal plate and deeply incised head, short, finely denticulate cusp and one pair of 
larger denticles. Laterals with broad heads and strong denticulate cusps, with 
two or more side denticles, and tapering, out-turned basal plates. The cusp lengthens 
as the distance from the rhachidian tooth increases, and becomes long, straight 
and spiky, losing its denticles except for a stout one laterally. The pattern is of a 
series of long straight cusps, each row overlapping the succeeding row. The four 
outermost teeth are degenerate. 

Caecum (Text-fig. 8b) large, lying flat on the surface of the digestive gland where 
exposed. 

Cerebral ganglia (Text-fig. 8a) fused, but may show their double origin. Visceral 
ganglia also fused. 

Specific Characters 

Large bulky Aplysias, of basic yellowish green colour, with numerous large black 
rings of irregular sizes on the sides of the body, thick rhinophores with short notched 
apices, broad foot with blunt tail, frilled swimming lobes joined low down posteriorly, 
broad spatulate spirally grooved penis, compound uniporous opaline gland, purple 
secretion, lateral teeth of the radula with long straight smooth cusps. 

Circum tropical. 

References [see also synonyms) 

Allan, 1932 : 420, as angasi ; 1950 : 212. 

Angas, 1867 : 228, as tigrina : 1877 : 190, as angasi. 

Arango, 1878 : 155, as protea. 

Baba, 1936 : 7 and 1949 : 124. 

Bergh, 1905 : 6, as benedicti. 

Dakin, Bennett & Pope, 1953 : 264, as angasi. 

Dall & Simpson, 1902 : 365, as T. protea. 

Dobson, 1880 : 159. 

Eales, 1944 : 3' &s fimbriata ; 1957 : I 79« 



310 REVISION OF THE WORLD SPECIES OF APLYS1A 

Engel, 1927 : 84 ; 1929 : 147 ; 1936 : 5. 

Gould, 1852 : 210, as fimbriata. 

Hedley, 19 10 : 370, as Tethys tigrina. 

MacFarland, 1909 : 14, as Tethys. 

Macnae, 1955 : 226 ; 1957 : 2 &9- 

Marcus E. & E., 1955 : J 4- 

Martens, 1880 : 307, as tigrina ; 1894 : 93. 

Morch, 1862 : 23, as protea ; 1875 : 176. 

Odhner, 1932 : 27. 

Olmsted, 1917 : 225. 

d'Orbigny, 1854 : 117, as protea. 

Pruvot-Fol, 1933 : 104 ; 1934 : 3 8 > IQ 47 : m ; 1953 " 33- 

Quoy & Gaimard, 1832 : 311. 

Rochebrune, 1881 : 264. 

Taki, 1932 : 212, as Tethys. 

Thiele, 1910:123, as Tethys. 

Winckworth, 1927 : 95, as benedicti. 

Aplysia (Varria) denisoni Smith, 1884 : 89 

Text-fig. 21 

Distribution. Indian Ocean (Cocos Keeling Islands), West Australia (Abrolhos 
Islands) ; South Pacific, Queensland and New South Wales. 

Material examined. Specimens from Cocos Keeling Island and Eastern 
Australia. 

The species was described by E. A. Smith from Queensland. Later O'Donoghue 
(1924 : 531) recorded and described it from the Abrolhos Islands off West Australia. 

Aplysias of moderate size, up to 150 mm. long, 32 mm. broad and 61 mm. high 
when preserved (Text-fig. 21a). Olive green, with dark mottlings and black veining. 
Inner sides of parapodia with vertical black and brown bands. No pigment on the 
foot, mantle roof and ctenidium, but this may have faded, and all trace of colour 
may disappear with long continued preservation. Skin soft, but not flabby, smooth. 

Head and neck large, broad and rather flat. Cephalic tentacles very wide, flat 
and plate-like, only slightly rolled on the edges (Text-fig. 21b). Eyes rather large. 
Rhinophores small, close together. 

Foot large, muscular but not tough, edges not well defined, tail short, pointed. 
Penis short, broad and spatulate. Parapodia small, low, commence far back, rounded, 
edges irregular but not fimbriated, joined low down on the foot, making a platform 
or low wall behind the visceral mass. Dorsal slit wide open. 

Mantle thin, weak, not large, with small conical papilla at the point of closure of 
the shell covering. Purple glands present. Anal siphon large, broad, jagged. 

Mantle cavity widely open, exposing the large ctenidium. Opaline gland compound, 
uniporous. Genital aperture plain, smooth, crescentic, the duct not inflated beneath 
the mantle cavity floor, genital groove weak. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



3«i 



Shell (Text-fig. 21c) ovate, broad, firm, with flat calcified apex, which is oblique. 
Anal sinus long and moderately deep. About ten fays spread from the apex to the 
anterior margin. 

Buccal mass small. Jaws narrow, rods very long, slender, slightly curved at the 
apex, loosely arranged. 




d) 



+ 1 



+6 +7 



A<&Wh 



Fig. 21. Aplysia denisoni. 




(a) Lateral view of a specimen from Australia, x i . 

(b) Head to show the large, plate-like cephalic tentacles. 

(c) Shell in dorsal and ventral views, x f . 

(d) Radular teeth. Rhachidian, first, sixth, seventh, eighth and outermost lateral 
teeth, x 8o. 



Radula (Text-fig. 2id) broad, 8*5 x 9 mm. in 83 mm. specimen. Formula 70 x 
53.1.53. Rhachidian with broad flat basal plate, paler than the laterals, cusp 
with faint denticulations and one pair of large denticles. First lateral with short, 
almost rectangular basal plate, weakly denticulate cusp and a large lateral basal 
denticle. The main cusp lengthens considerably as distance from the centre increases, 
but the laterals remain simple, with smooth or feebly denticulate cusps and large 
basal denticles. Older teeth become quite smooth. Up to five vestigial teeth at 
the end of the row. 

Caecum small, appearing flat on the surface of the digestive gland. 

Cerebral ganglia fused as a flat band. Visceral ganglia joined but recognizably 
paired. 



3X2 . REVISION OF THE WORLD SPECIES OF APLYSIA 

Specific Characters 

Moderate size, very large plate-like cephalic tentacles, rounded parapodia joined 
low down on the foot posteriorly, minute mantle papilla, compound uniporous 
opaline gland, ovate calcified shell with narrow oblique apex and long anal sinus, 
broad radula with weak denticulations and elongated cusps to the laterals. 

Australia and southern Indian Ocean. 

References 
Hedley, 1909 : 370. 
Pilsbry, 1895 : 102, as Tethys. 

Aplysia (Varria) extraor dinar ia Allan, 1932 : 314 

Text-figs. 22 and 23 

Distribution. South Pacific, from Hawaiian Islands to New South Wales, 
Australia. 

Material examined. The type specimen from the Australian Museum, Sydney, 
taken by Miss Allan at Port Hacking, New South Wales, two specimens brought 
to this country for the International Fisheries Exhibition in 1883 from Port Jackson, 
now in the British Museum (Natural History), and one specimen from the Paris 
National Museum, collected by M. Ballieu in the Hawaiian Islands in 1874. 

A well defined species, large (at least 300 mm. long when alive), slender and active. 
It shrinks to about half this length when preserved, and while remaining soft and 
flabby may become intensely wrinkled. The type specimen (Text-fig. 22) measures 
146 mm. long, 32 mm. wide and 73 mm. high. The colour is deep umber brown 
with reticulate black veining and scattered black or brown blotches. There may 
also be light spots and vertical white bands, and black spots on the head. The inner 
sides of the parapodia have vertical bands of light and dark near the edges, but oval 
light areas on a black background below. The mantle is mottled brown and white, 
with concentric lines of brown pigment along the edges. All pigment may bleach 
as the result of long continued preservation. 

Head small, long and narrow, with slender neck (Text-fig. 23a). Cephalic tentacles 
wide, continued as broad rounded side flaps to the mouth. Eyes small. Rhinophores 
acutely pointed, slender, very close together, set far back and slit more than half-way. 

Foot very long and narrow, only 22-26 mm. wide in the type, soft and not thick. 
Anterior border rounded and slightly enlarged, edges not well defined laterally, 
tail long and slender. Traces of pigment remain. Penis sheath small, penis elongated, 
stout at the base, pointed, with straight sperm groove to the tip (Text-fig. 23c). 
Parapodia large, rounded, arising close together and meeting low down posteriorly, 
forming a 24 mm. extension of the mantle cavity floor. In another specimen they 
appear to be free. Edges sinuous or smooth. Mobile and probably natatory. 



REVISION OF THE WORLD SPECIES OF APLYSIA 313 

Mantle small, 55 x 30 mm., thin, with little overhang anteriorly, but a wide 
flap laterally. Foramen small, with contraction rings around it. Anal siphon 19 x 16 
mm. wide, tall, thin and leaf-like. Mantle glands well developed, spotted with dark 
pigment, so probably purple is secreted. 

Mantle cavity open at both ends. Ctenidium large, 30 x 15 mm. Opaline gland 




Fig. 22. Aplysia extraordinaria. Dorsal view of the type specimen. The sinuous edge 
of the parapodium is shown on the right, the pattern of the inner side on the left, with 
its light rim. The right mantle edge has been reflected to show the ctenidium and 
the purple gland. The small papilla on the mantle is ringed, and the mantle is 
blotched with brown and white, but is lined near the edge. X f . An enlarged 
view of the mouth region is alongside. 

small, simple, multiporous. Genital aperture not pronounced, plain edged. Genital 
groove slight, marked by cross stripes of brown in the type. 

Shell (Text-fig. 236) removed from the type. Said by Allan to be 75 X 56 mm. 
In another specimen it measured 45 x 36 mm., and is therefore broadly ovate, 
with strong concentric lines, without spire, anal sinus shallow. Deep yellow in 
colour. 

The jaws are curved plates, composed of closely packed rods, rather short, trun- 
cate at the apex. 

ZOOL. 5, IO. l6 



314 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Radula (Text-fig. 23d) with about 70 rows and a formula 32.1.32. Rhachidian 
tooth pale, weak and sunken, with a short and not broad basal plate, almost straight 
posteriorly and with a short cusp with fine denticulations. The first lateral has 




Fig. 23. Aplysia extraordinaria. 

(a) Half lateral view of a specimen in the B.M. (N.H.) collection to show the elongation 
of the neck region, the large parapodia and the tall, leaf-like anal siphon. The head 
is somewhat contracted, x f . 

(b) Shell in dorsal view, x f . 

(c) Penis sheath with its retractor muscles, and the extruded penis. X 1 J. 

(d) Radular teeth. Rhachidian, first, sixth and 20th lateral teeth X 80, the last 
enlarged, x 140. 



a narrow basal plate enlarging to a swollen head, short cusp and numerous denticles 
on both sides, the lateral basal denticle larger than the others and its edge cut into 
denticles ; this feature is repeated throughout, the denticulations becoming highly 
elaborate and irregular, though the cusps remain of moderate length. The three 
outermost teeth are vestigial. 



REVISION OF THE WORLD SPECIES OF APLYSIA 315 

Caecum small, lying flat on the surface of the digestive gland. In the type specimen 
it is corrugated and its tip is recurved, but in another specimen it remains straight. 
Cerebral ganglia fused, visceral joined and spread along the visceral cords. 

Specific Characters 

Large Aplysias, not bulky but slender and active. Deep brown with black reticu- 
lations and scattered spots and blotches. Skin smooth, soft and flabby. Head small 
with slender neck. Rhinophores slender, close together. Foot long and narrow, 
tail long. Parapodia large, rounded, natatory, meeting low down posteriorly. 
Mantle small, with contractile foramen and tall anal siphon. Purple secretion 
probable. Opaline gland simple, small, multiporous. Shell broadly ovate. Radula 
with elaborate primary and secondary denticulations. 

South Pacific area. 

Aplysia (Varria) fasciata Poiret, 1789 : 2 

Text-fig. 24 

Synonymy : alba Cuvier, 1803 : 295 ; Rang, 1828 : 60, pi. 15. 
cameliformis Locard, 1886 : 66. 
camelus Cuvier, 1803 : 295 ; Rang, 1828 : 60, pi. 15. 
depilans ; de Blainville, 1823 : 286. 
leporina Rang, 1828 : 54, pis. 6 and 7. 
lepus Philippi, 1844 : 99. 

limacina de Blainville, 18236 : 287 ; a : 328 ; Blochmann, 1884 : 29 
marmorata de Blainville, 1823& : 286 ; a : 326. 
neapolitana and napolitana delle Chiaje, 1823 : 31, 39, 70. 
radiata Crouch, 1826 : 44. 
sicula? Swainson 1840 : 247, fig. 45. 
vulgaris de Blainville, 1823 : 285. 
Dolabella lepus Risso, 1826 : 44. 
Lepus marinus Rondelet, 1554 : 520. 
Lernaea Bohadsch, 1761 : 49. Not binominal, but two kinds are 

recognized, one of which is fasciata (see p. 357). 
Siphonota lobiancoi Mazzarelli, 1890 : 42 (probably a hybrid). 

Distribution. Red Sea, Mediterranean, Atlantic coasts from France to Angola 
(Portuguese West Africa), taken in Morocco, Senegal, Ghana and the Canary Islands. 
Rarely occurs on the southern coasts of Britain. 

Material examined. Specimens from the above areas. 

This handsome species was described by Poiret from the coast of Barbary in its 
black, scarlet-bordered form. Rang (1828, pis. 6 and 7) gave good coloured figures of 



3 i6 REVISION OF THE WORLD SPECIES OF APLYSIA 

it, and on pi. 15 and 15 bis. as camelus, alba and napolitana. Several authors confused 
it with depilans. Pilsbry reverted to Linnaeus's name leporina, because, he says, 
" Rondelet's figure and description of the coloration agree well with this species 
and not with any other European Sea Hare ". The International Commission on 
Zoological Nomenclature, however, (Opinion 200, 1954 and Opinion 354, 1955) 
validated the name fasciata Poiret, 1789, owing to confusion between leporina, 
depilans and the nudibranch Tethys leporina (now fimbria) . 

Large Aplysias, reaching nearly 400 mm. when alive. High and narrow, a pre- 
served specimen measuring 138 mm. long, 60 mm. wide and 90 mm. high. Skin 




Fig. 24. Aplysia fasciata. 

(a) Penis sheath opened longitudinally to expose the long, filiform penis. The seminal 
groove is indicated but not continued to the tip of the penis owing to the narrowness 
of the latter. A small flap lies at its base, x 2. 

(b) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth, x 80. 



soft and firm. Colour velvety black all over, or spotted with clusters of white and 
grey flecks and with a red border to the parapodia, cephalic tentacles and rhinophores, 
fading to a light rim with preservation. The black colour is marked on the mantle, 
ctenidium and anal siphon, but the mantle cavity floor and foot are usually less 
heavily pigmented. Paler specimens may be yellow, with round brown or black 
spots, and occasionally white individuals occur. All colour may disappear with 
long preservation. 

Head large, neck short owing to the forward position of the parapodia, but capable 
of considerable extension. Cephalic tentacles well developed but not long, rolled 
at the edges, fimbriated on the unusually large rounded extensions to the ventral 
sides of the mouth slit. Eyes small. Rhinophores slender, small, acutely pointed, 



REVISION OF THE WORLD SPECIES OF APLYSIA 317 

slit for only a short distance, the rims of the slit crimson or pale, set close together, 
only about 9 mm. apart. 

Foot narrow, elongated, the lateral edges approximating towards one another 
as in punctata. Front edges rounded, tail short, pointed. Penis (Text-fig. 24a) 
long and slender, filiform, unpigmented. Rang (1828, pi. 7) gave a good figure of it. 
Distal (bulbous) portion of the sheath small, smooth-lined, with a flap-like ingrowth 
alongside the sperm groove but not touching it. Parapodia very large, mobile, 
natatory, arising close together about 20 mm. apart, rounded, tall, with smooth 
borders, meeting low down on the tail and so narrow here that they are sometimes 
described as free. The mantle cavity is thus open posteriorly. The parapodia will 
close completely over the mantle, leaving the anal siphon projecting. 

Mantle large, foramen a small flat hole, usually rayed, and tubular if much con- 
tracted. Purple secretion exuded. Anal siphon large, broad, tubular. 

Mantle cavity widely open. Ctenidium large and thick. Opaline gland compound, 
compact, with a small round aperture. Genital aperture oblique, fimbriated. 

Shell about 70 x 50 mm., broadly ovate, concave, firm, without spire, anal sinus 
short and not deep. 

Jaws rectangular, rods elongate, curved at the tips. 

Radula (Text-fig. 246) large, reaching a formula of 80 x 50.1.50. Rhachidian 
tooth of the same colour as the laterals, not pale or sunken, its basal plate of moderate 
size, the head tall and narrow, the cusp longer than the plate, rounded, with numerous 
weak, irregular denticles, the basal ones only slightly larger than the others. First 
lateral with curved tapering plate, moderately broad head and the usual denticles, 
a lateral basal one enlarged. Along the row the plates remain tall and narrow, the 
cusps lengthen but rarely exceed more than half the length of the plate and the denti- 
cles become stronger, often with secondary denticulations of the large lateral basal 
denticles and the addition of others near the head. About four teeth at the end of 
the row are vestigial. 

Caecum tip visible on the surface of the digestive gland. 

Cerebral ganglia small, fused. 

Specific Characters 

Large, narrow and high Aplysias, typically black with scarlet rims to the parapodia 
and tentacles. Head large, with fimbriated cephalic tentacles, narrow foot with 
pointed tail, penis filiform, white, parapodia large, natatory, meeting low down 
on the tail, mantle with small flat foramen, purple secretion, compound uniporous 
opaline gland, shell broadly ovate, radula large, all the teeth denticulate. 

East Atlantic, from France to Angola ; Mediterranean and Red Seas. 

References {see also synonyms) 
Bosc, 1802 : 63, as Laplisia. 
Cantraine, 1840 : 68. 
Cams, 1889 : 196. 
Eales, 1957 : 179, as leporina. 
Engel, 1934 : 85, 88. 



318 REVISION OF THE WORLD SPECIES OF THE 

Grigg, 1949 : 796, as limacina. 

Mazzarelli, 1893 : 33, as Siphonota limacina. 

Odhner, 1932 : 27, as leporina. 

Pilsbry, 1895 : 72, as Tethys leporina. 

Rang, 1828 : 54, 60, pi. 15 (as alba, camelus). 

Risbec, 1931 : 68. 

Vayssiere, 1885 : 54, 60 ; 1935 : No. 20 (not paginated). 

Verany, 1853 : 390. 

Aplysia (Varria) gigantea Sowerby, 1869, pi. 1 
Text-fig. 25 
Synonymy : A. tasmanica Tenison Woods, 1876 : 156. 

Distribution. Australian region — West Australia, Abrolhos Islands, New 
South Wales, Tasmania. 

Sowerby named the species from the shell. O'Donoghue (1924) described specimens 
from near Fremantle, Australia, and these are now in the British Museum (Natural 
History) . 

Material examined. O'Donoghue's topotypes (B.M. (N.H.)), 1923. 1.26.10-11. 

Aplysias of enormous size, reaching 600 mm. in length, but shrinking to much less 
when preserved. 150 mm. long x 65 mm. wide X 60 mm. high is an average preserved 
specimen. Body stout and high. Colour and markings variable, background colour 
dark umber brown to purplish green, with black spots, branched black lines and 
blotches of light, almost white, areas. The edges of the parapodia may be banded 
vertically, both inside and out, with alternating bands of white and dark, together 
with similar blotches of lighter colour, and there is a fine reticulation extending to 
the edge externally (Text-fig. 25, b, c). Foot greyish black. Mantle floor and ctenidium 
usually unpigmented or slightly pigmented. On contracting, the soft fleshy skin 
assumes a crocodile-skin pattern of flattened areas interrupted by furrows. This is 
exhibited by small as well as by full grown specimens, and is characteristic though 
not unique. 

Head and neck long. Cephalic tentacles very large, rolled laterally and continued 
to the sides of the mouth. Rhinophores long and linear, set close together. 

Foot broad, especially anteriorly, where it is expanded at the lateral borders 
and measures 85 mm. wide in a specimen only 150 mm. long (Text-fig. 25a). The 
tail has similar but smaller expansions. Edges of the foot wrinkled, foot sole very 
muscular, contracting to the crocodile-skin pattern. Penis very broad and stout, 
short and flat. Parapodia high and well developed, mobile and natatory, rounded 
in shape, with thin irregular edges. They commence far forward and stand straight 
up. Posteriorly they are united low down on the foot, forming a spout-like projection 
of the mantle cavity floor. The dorsal slit is wide open, exposing both mantle and 
ctenidium. 

Mantle large, thin, perforated by a minute foramen on a low papilla, sometimes 
wrinkled radially. Anal siphon large and broad. Purple glands present. 



REVISION OF THE WORPD SPECIES OF APLYSIA 



319 



Mantle cavity widely open, large. Ctenidium very large and protruding. Opaline 
gland compound with one aperture. Genital aperture smooth or slightly fimbriated. 

Shell large. Sowerby figures a dried specimen measuring 96 x 79 mm. It is 
thin, transparent, pale yellow, concave, the apex a little incurved, the anal sinus 
long and shallow. Calcareous matter is present during life. 

Jaws oblique, composed of slender, curved rods of moderate length. 

Radula 15-5 X 14-5 mm., very strong. O'Donoghue (1924) gives 85 rows and 
formula 59. 1 .59. Rhachidian with short broad basal plate, very short cusp bearing 




Fig. 25. Aplysia gigantea. 

(a) Head and foot in ventral view to show the large lateral wings on the foot. x 

(b) Portion of the outer side of the parapodium to show the fine reticulations. 

(c) Inner side of the parapodium, with branched bands of black on a light ground. 



feeble denticles. First lateral with short plate and cusp, but very wide head, its 
denticles feeble, but of the usual pattern. Outer laterals may become smooth except 
for one side denticle on a straight slender cusp. The basal plates of the laterals are 
squared off and packed in close lines, but the last six or eight teeth are more slender, 
the outermost three being vestigial. 

The inflated tip of the caecum just reaches the surface of the digestive gland. 

Cerebral ganglia completely fused, and also the visceral pair. 



Specific Characters 

Large size, smooth skin contracting to a crocodile-skin pattern, large cephalic 
tentacles, foot expanded laterally anteriorly, high rounded parapodia joined low 
down on the foot posteriorly, minute mantle aperture and papilla, compound 



320 REVISION OF THE WORLD SPECIES OF APLYSIA 

uniporous opaline gland, large rather shapeless shell, numerous closely packed 
radular teeth with feeble denticulations. 
Australia and islands off its coasts. 



References [see also synonym) 



Allan, 1950 1213. 

O'Donoghue, 1924 : 528. as Tethys. 

Pilsbry, 1895 : 102, as Tethys. 

Aplysia (Varria) gracilis sp. nov. 

Text-fig. 26 
Distribution. Red Sea. 

Material examined. A single specimen, the type, B.M. (N.H.), 1907. n. 15. 29), 
presented by His Excellency the Minister of Education to the Government of Egypt, 
and collected near Gemil, Lake Manzaleh in 1907. 

A small Aplysia, the preserved specimen measuring 45 mm. long x 25 mm. wide 
X 21 mm. high, much contracted to the " sitting hare " position. Skin soft, mottled 
with brownish black and grey on a lighter greenish grey background. There are 
a few solid spots on the sides. The mantle has blotches of pigment and there are 
faint markings inside the parapodia, mostly vertical bands of light and dark colour. 
The tentacles, mantle cavity floor, ctenidium and foot are colourless. The rims of 
the parapodia and anal siphon are light and devoid of pigment. 

Head and neck small. Cephalic tentacles rolled and continued forwards to the lower 
sides of the mouth. Eyes small but plainly visible. Rhinophores close together and 
slit almost to their bases, small and conical. 

Foot narrow, not tough, 43 mm. long and only 8 mm. wide anteriorly, rounded at 
the front edges, furrowed longitudinally and prolonged into a short pointed tail. 
It is sharply marked off from the pigmented side of the body by a small flange. 
Penis slender and tapering but not very long, the sheath small and smooth-lined 
distally. Parapodia fairly large and thick but not high, arising far forwards on the 
neck, sloping backwards, with pale sinuous fimbriated edges, to meet low down on 
the foot posteriorly, so that the mantle cavity is open and flat. 

Mantle small and thin, measuring 20 x 11 mm. There is a small aperture on a 
papilla. Anal siphon wide and high, directed towards the opening in the parapodia 
(i.e. backwards) in the specimen. Mantle glands well developed, presumably secret- 
ing purple. 

Mantle cavity small, ctenidium small and neat. Opaline gland compound, with 
a single large circular aperture. Genital aperture thin, oblique, pigmented, deeply 
crescentic. Genital groove shallow and colourless. 

Shell small, ovate, delicate, measuring 18 X 14 mm., rather flat. Calcareous lining 
present. There is hardly any turn over at the apex and the anal sinus is shallow. 

Jaws pale and delicate, the rods moderately tall, broadened a little at the tips 
and slightly curved. 



REVISION OF THE WORLD SPECIES OF APLYSIA 321 

Radula not large, almost square, 4x4 mm., with a formula 40 x 21.1.21. 
It is the most elaborate of all the species of the genus (Text-fig. 26) . Rhachidian 
tooth with a short broad basal plate, slightly incised on its posterior border, but 
becoming more hollowed out in older teeth. Cusp narrow, at first rounded, then 
acuminate, longer than the plate, with numerous, short, irregular denticles of which 
the basal pair are as usual larger than the others. The first lateral has a narrow 



4-6 





Fig. 26. Aplysia gracilis. Radular teeth. Rhachidian, first, sixth and five outermost 
lateral teeth. x 80. Eighth lateral enlarged. x 132. 

out-turned plate, large head and more regular denticles, the two outer basal ones 
being larger than the others and the penultimate one very stout. The cusps increase 
in length up to about the middle of the row, and their denticles become very large 
and deeply cut. Mesially there are two small denticles and one larger basal denticle, 
laterally up to nine denticles, the longest and broadest being the seventh. The last 
four teeth are degenerate. 

Caecum exposed and lying flat on the surface of the digestive gland. 

Cerebral ganglia joined, visceral cords short and the ganglia joined but distinct. 

Specific Characters 

Everything about this species is neat and dainty. The specimen is probably not 
full grown. Small, with mottled and spotted skin, narrow foot, slender pointed 
penis, fimbriated parapodia joined low down behind, minute papilla on the mantle, 
compound uniporous opaline gland, delicate oval shell with feeble apex and anal 
sinus, highly elaborate radula with deeply cut denticulations. 

Red Sea. 

Aplysia (Varria) inca d'Orbigny, 1837 : 20 7 
Distribution. Recorded from Callao Bay, near Lima, Peru. 

Material examined. The type (1834) an d a second specimen collected by Eydoux 
& Souleyet in 1838, now in the Museum d'Histoire naturelle, Paris, were examined. 
Good coloured figures were given by d'Orbigny and reproduced by Pilsbry (1895). 
Both the specimens in the Paris Museum, however, have lost their colour and are 
much reduced in size ; neither has a radula, the type has had the shell removed, 
and the second specimen is without the penis. 



322 REVISION OF THE WORLD SPECIES OF APLYSIA 

The type specimen now measures 60 mm. long, 17 mm. wide and 26 mm. high, 
the small mantle is 24 X 13 mm., the foot 54 X 13 mm. D'Orbigny described the 
living animal as large (200 mm.), flabby and swollen. The cephalic tentacles in his 
figure are very wide and rolled, the rhinophores set well back and close together, 
conical, slit. Eyes plain. Neck shorter than the distance between the cephalic 
tentacles and the rhinophores. Foot narrow, with pointed tail. Parapodia large, 
united posteriorly. Mantle smooth, with a small round aperture and long, wide and 
thin anal siphon. Animal violet, with round white spots on the sides and in a row on 
each side of the head. Inner sides of the parapodia with a narrow border of clear 
rose violet and large rounded and angular white blotches on a purple brown ground. 
Mantle violet. 

To this may be added : Penis short, broad, tapering and flat at the tip. Penis 
sheath smooth-lined distally. Parapodia smooth edged, joined to form a low wall 
about 10 mm. high around the mantle cavity posteriorly. 

Opaline gland small, partly compound, but with a group of apertures arranged in 
a circle. 

Shell broadly ovate with distinct anal sinus. 

Caecum small, just breaking surface on the digestive gland. 

Cerebral ganglia joined, visceral ganglia distinct. 

Specific Characters 

Large size, purple coloration with white spots, soft flabby skin, capable of great 
contraction, wide cephalic tentacles, short broad tapering penis, narrow foot and 
pointed tail, large parapodia united to form a low wall posteriorly, small mantle 
aperture, partly compound opaline gland with a group of apertures. Radula unknown. 

Eastern South Pacific. 

Reference 
Pilsbry, 1895 : 87, as Tethys. 

Aplysia (Varria) keraudreni Rang, 1828 : 59, pi. 13 
Text-fig. 27 

Synonymy : lessoni? Rang, 1828 : 60, pi. 14. See below. 

Distribution. The species has been recorded from the South Pacific only. Rang's 
single specimen came from the Society Islands. 

Material examined. Five specimens from the Portobello Marine Biological 
Station, Otago, New Zealand, and one from Wellington Harbour ; another, juvenile, 
from the Kermadec Islands. A small specimen, much contracted, but probably 
belonging to this species, came from Sydney, Australia. 

Rang gave a good figure of this large and handsome species, copied by Pilsbry 

(1895: pi. 39). 



REVISION OF THE WORLD SPECIES OF APLYSIA 



3*3 



Mature specimens attain 250 mm. or more in length, preserved about 188 mm. 
long, 75 mm. wide and 80 mm. high. Thick and fleshy, with rather small head, but 
very large mantle region and visceral hump. Olive green to dark brownish black, 
with scattered brown spots or blotches and reticulate black lines all over. There 
may also be white areas. Cephalic tentacles, rhinophores and edges of the parapodia 
with light brown or white margins. Mantle dappled light and dark. Inner sides 
of the parapodia with irregular light flecks on a dark ground. Skin soft, but when 



sem.gr 




Fig. 27. Apiysia keraudreni. 

(a) The extruded penis, showing its filiform shape and the two portions of the sheath, the 
distal part of which is white and fibrous, the proximal part thick, dark and muscular. 
The seminal groove has been sketched as far as the penis base only, x 2. 

(b) Shell in dorsal and ventral views. X f . 

(c) Radular teeth. Rhachidian, first, sixth, 12th and outermost lateral teeth, x 80. 



contracted may form crocodile-skin patterns or become firm and tough. All colour 
may disappear with preservation. 

Head rather small but broad, neck short, thick. Cephalic tentacles large, rolled 
on the edges, rounded in front, joined to the sides of the mouth slit, fimbriated. 
Eyes small. Rhinophores slender, pointed, close together, slit half-way and dark 
within the slit. 

Foot broad, 50 mm. wide in the 188 mm. specimen, thick, spongy, prolonged into 
a bluntly pointed tail about 28 mm. long. Penis and its sheath characteristic and 
easily eversible. Penis (Text-fig. 27a) white, filiform, the distal bulbous portion of 
the sheath also white, thin-walled, the proximal muscular portion dark, rugose. 
Parapodia large, rounded, frilled on the edges or smooth according to the degree 



3 2 4 REVISION OF THE WORLD SPECIES OF APLYSIA 

of contraction. Only 23 mm. apart in front, rather thick, freely mobile, natatory, 
meeting low down and hardly joining on the root of the tail, forming a flat platform, 
or a low wall about 10 mm. high around the mantle cavity. 

Mantle very large, 70 x 52 mm., thick, with small, round, finely rayed aperture. 
Anal siphon broad, frilled but not tall, tubular. Mantle glands well developed, 
exuding purple. Mantle border wide, overlapping the ctenidium. 

Mantle cavity large, ctenidium also large, with some pigment. Opaline gland 
simple, with large vesicles and numerous apertures, the gland cells tending to join 
anteriorly by fusion of their ducts and forming an anterior aperture larger than the 
others. Genital aperture small, smooth, not covered by the mantle or pigmented. 

Shell (Text-fig. 276), large, 70 x 50 mm., broadly oval, shallow, with weak apex 
just recurved and dorsal border bent over. Anal sinus short, thick walled, shallow, 
with rounded outer angle. Lines of growth well marked. 

Jaws long and rectangular, 10 X 2-5 mm., rods slender, curved or straight, of 
moderate length. 

Radula 13 x 10 mm., with a formula 55 x 35. 1.35 (Text-fig. 27c). Rhachidian 
tooth pale, sunken, with small and rather narrow basal plate, excavated in front, 
straight behind. The cusp is equal to or a little longer than the plate, rounded, 
with irregular denticles, fluted, the basal denticles larger than the others. First 
lateral with broad, elevated head, similar denticles laterally and short rounded cusp, 
also fluted. The remaining teeth with broad low heads, lengthening and narrowing 
cusps and great exaggeration of the lateral denticles in number, size and irregularity. 
Secondary denticulations occur in some. The outermost three to four teeth rectangu- 
lar and vestigial. A very elaborate radula. 

Caecum just appearing on the surface, slightly bent at the tip. 

Cerebral ganglia completely fused, visceral with large nerve cells making them 
appear knobbly. 

Specific Characters 

Large handsome Aplysias, with fimbriated cephalic tentacles and parapodia. 
Colour dark, with spots and blotches of a darker colour, but with light edges to 
the tentacles and parapodia. Head rather small, visceral region exceptionally 
large, with wide overhang to the mantle, where the purple glands lie. Foot broad, 
with blunt tail. Penis white, filiform, penis sheath partly white thin, partly dark 
muscular. Parapodia large, natatory, meeting low on the tail. Mantle aperture 
small, rayed. Opaline gland simple, multiporous. Shell large, broadly ovate, shallow, 
apex narrowed. Radula elaborately denticulate. 

South Pacific. 

References [see also synonym) 
Lesson, 1830 : 294. 
Mazzarelli & Zuccardi, 1892 : 11. 
Pilsbry, 1895 : 95, as Tethys. 

Note. Angas's Syphonota keraudreni, 1867 and Sowerby's A. keraudreni from 
Australia are probably A. dactylomela. 



REVISION OF THE WORLD SPECIES OF APLYSIA 325 

I have not seen A. lessoni Rang, 1828 : 60, recorded from Peru, and by Mazzarelli 
& Zuccardi (1892 : 11) from Honolulu. From the little that is known of it, I suspect 
that it is keraudreni, with which it agrees in size, shape, large, rounded cephalic 
tentacles, broad foot, large parapodia united low behind, large mantle with tubular 
aperture and shape of the shell. Mazzarelli & Zuccardi added the information that 
the opaline gland is simple, multiporous, the radula with 30 rows and formula 
45.1.45. Their figure of the radula is similar to that found in keraudreni. 



Aplysia (Varria) kurodai Baba, 1937 : 213 

Synonymy : Takahashi's Tethys punctata Cuvier, from Formosa (1934 : 357) is 
probably this species. 

Distribution. Japan, Formosa, Asiatic Mainland. Baba states that it is a 
common species in Japan and Formosa. 

Material examined. Seven good specimens, hitherto unnamed, have been in 
the British Museum (Nat. Hist.) collections since 1921 and were collected by A. V. 
Insole at Misaki, Japan. The Paris National Museum has 18 specimens from Japan, 
collected in 1903, and two from the mainland of Manchukuo, dated 1844. 

Baba (1949 : pi. 14) gave coloured figures of the species, and a sketch of the radula 
(p. 25). In 1956 he described the egg strings as pink or yellow, with 15-30 eggs in 
each capsule. 

Large Aplysias, attaining 400 mm. in length. Colour variable, but typically dark, 
brown or purplish black, with small irregular greyish mottling on the sides and mantle, 
clear blotches on the inner sides of the parapodia and sometimes black edges. Foot 
dark, mantle cavity floor and ctenidium not pigmented. The general appearance is 
that of a very heavily pigmented animal. Skin soft, but not flabby. On contracting 
the species assumes the sitting hare position. Baba's figure shows it to be a mobile 
species. 

Head and neck long but not large. Cephalic tentacles more than double the length 
of the rhinophores, sinuous on the edges, folded. Rhinophores narrow, pointed, 
but stout when contracted. 

Foot large, broad, 22 mm. wide in a 62 mm. specimen, tough, truncate anteriorly, 
much wrinkled in contraction, tail short, obtuse. Penis and sheath small, penis 
filiform. Genital groove delicate. Parapodia rounded, thin, flexible, black and 
irregular on the edges, joined low down on the tail, often appearing to be quite free. 
They hardly cover the mantle. 

Mantle oval, rather small ; a small foramen opens on a papilla, but is not easily 
seen owing to the heavy pigmentation and mottled pattern of this area. Anal 
siphon short, broad, tubular ; anus with lobed aperture. Purple glands present. 

Mantle cavity open. Ctenidium small, opaline gland kidney shaped, simple, 
multiporous. Genital aperture small, crescentic, smooth. 



326 REVISION OF THE WORLD SPECIES OF APLYSIA 

Shell oval, narrow (30 x 19 mm. in a 62 mm. specimen), rather flat, thin ; apical 
region weak, but the edge recurved and thickened dorsally ; anal sinus shallow, 
calcareous matter present. 

Jaws dark brown, rods long and curved. 

Radula 50 rows, 34.1.34, but Baba gave 70 x 43.1.43. Teeth closely packed. 
Rhachidian tooth with moderately broad basal plate, incised anteriorly but almost 
straight posteriorly, with rounded edges. Cusp hardly longer than the plate, but 
becoming shorter with wear. Denticles irregular. First lateral with broad curved 
out-turned basal plate, large head and irregular denticles on the cusp, with two 
larger basal denticles laterally. Sixth lateral with broad rounded basal plate, a 
longer and sharper cusp, but with similar irregularities in the denticulation. Last 
three teeth vestigial. 

Caecum very large, curved, swollen at the tip. 

Cerebral ganglia fused, visceral fused. 

Specific Characters 

Large Aplysias, with heavy pigmentation, mobile. Penis filiform ; foot broad ; 
mantle papilla small ; mantle glands secreting purple ; opaline gland simple multi- 
porous ; shell flat, thin, radula large with numerous teeth, irregular denticles ; 
chief ganglia fused. 

North-west Pacific. 

Reference 

Baba, Hamatani & Hisai, 1956 : 216 (breeding habits). 



Aplysia (Varria) maculata Rang, 1828 : 58, pi. 12 

Synonymy : eusiphonata Bergh, 1908 : 9. 
gargantua Bergh, 1908 : 5. 
gilchristi Bergh, 1908 : 2. 
poikilia Bergh, 1908 : 10. 
Tethys poikilia O'Donoghue, 1929 : 21. 
tigrina ; Quoy & Gaimard, 1832 : 308 (non Rang, 1828). 
iigrinella Gray, M.E., 1850 : 97, new name for A. tigrina Q. & G. 

Distribution. Confined to the western part of the Indian Ocean : Mauritius, 
Reunion, Lourenco Marques, South Africa. 

Specimens examined. Several from South Africa, sent by Dr. Macnae. 

Rang gave good figures of this species, copied by Pilsbry (1895 : pi. 17). In 1955 
Macnae published a detailed account of its structure and a useful compilation of the 
synonyms. He decided that Quoy & Gaimard' s tigrina, renamed tigrinella by Mrs. 
Gray, and re-examined by Mme. Pruvot-Fol (1934 : 39), is a synonym of maculata, 
thus clearing up the confusion arising from Quoy & Gaimard's misuse of Rang's 



REVISION OF THE WORLD SPECIES OF APLYSIA 327 

name. The papillae shown in their figure (see Pilsbry, 1895 : pi. 16) were not men- 
tioned in the description and may have been added mistakenly by the engraver. 
Papillae, however, do occasionally occur in preserved specimens of other species, 
cf. A. parvula above (p. 288). 

Rather large Aplysias, with long neck and ovate posterior region, ending in a 
narrow pointed tail. May attain 300 mm. in length, but a preserved specimen 
shrinks to about 170 mm. long, 34 mm. wide and 77 mm. high. The skin is soft and 
flabby. Colour very variable, usually brownish olive, with irregular black and light 
spots, often confluent in patches on the head, sides of the body and inner sides of the 
parapodia. A few spots on the foot and mantle, the latter sometimes with irregular 
radiating lines of black. Some specimens are grey all over, without black pigment, 
or some of the spots may have clear centres and become rings, but this is not common. 
The parapodial rims are unpigmented or white. 

Head and neck narrow and long, usually narrowed behind the head. Cephalic 
tentacles well developed, capable of considerable expansion, not fimbriated, rolled on 
the edges and continued to the borders of the mouth. Eyes small. Rhinophores 
slender, tapering, close together and well back. 

Foot narrow (only 20 mm. broad in a specimen 85 mm. long). Rounded anteriorly, 
prolonged posteriorly into a pointed tail. Lateral edges well defined and usually 
projecting as a flange all round. Penis short, broad at the base, but tapering to a 
slender point, the sperm groove running straight to its apex. Parapodia large, 
extending from close behind the rhinophores to the root of the tail, thin, tall 
and wing-like, flexible but not fimbriated. Macnae states that the young animal 
can swim. Anteriorly the parapodia slope backwards, as Rang's figure showed ; 
posteriorly they join low down, or may appear free. The mantle cavity is thus 
open at both ends, the dorsal slit is wide open, but may be closed by folding over 
of the parapodia. 

Mantle large and thin. Mantle foramen small, usually flat, with radiating black 
lines around it. Anal siphon large, tubular, leaf-like, with a sinuous edge ; it has 
a tendency to slope backwards, as in many species in which the parapodia meet 
low down on the foot. Purple glands are present. 

Mantle cavity large and flat, the ctenidium large and unpigmented. Opaline 
gland simple and pear-shaped, multiporous. Genital aperture anterior to the mantle, 
plain and crescentic. 

Shell elongated (measuring 34 mm. x 22 mm. x 4 mm. in a 67 mm. specimen), 
thin and shallow, without a spire, the apex being only slightly recurved. Calcareous 
matter is present in the fresh condition. The anal sinus is short and shallow. 

Jaw plates well developed, dark brown in colour. The rods long and curved, with 
pointed or truncated tips. 

Buccal mass relatively large. Radula with about 65 rows and a formula up to 
38.1.38. The rhachidian tooth is of moderate height and breadth, its basal plate 
excavated in front and almost straight behind, the cusp about as long as the plate, 
with deeply cut denticles, the basal pair of which are larger than the others. Older 
teeth may become fluted and lose their denticles. The first lateral has a short cusp 



328 REVISION OF THE WORLD SPECIES OF APLYSIA 

and the usual denticles. Both the main cusp and the basal denticle become accentu- 
ated as the distance from the centre of the row increases, a strong denticle is added 
mesially and one or more laterally. As the cusp narrows it becomes curved lateral- 
wards, but retains its deeply cut denticles. After about the 24th tooth the cusp 
shortens, and the outermost six teeth are degenerate, a rather large number for an 
Aplysia. 

Caecum large and inflated at the tip, its blind end exposed on the surface for some 
distance. 

Cerebral ganglia completely fused, forming a flat band. Visceral ganglia joined 
but distinguishable. 

Specific Characters 

Small head, long neck, ovate visceral region, widely open dorsal slit, spotted skin, 
narrow foot with lateral flange and pointed tail, short acuminate penis, mobile 
parapodia joined or free low down posteriorly, large mantle with small rayed aperture, 
simple opaline gland with many apertures, delicate narrow shell and radula with 
deeply cut denticulations. 

Western Indian Ocean, from Mauritius to the Cape. 

References {see also synonyms) 
Deshayes, 1863 : 54. 
Krauss, 1848 : 71. 
Lamarck, 1836 : 691. 
Macnae, 1955 : 231. 
Martens, 1880 : 307. 
Pilsbry, 1895 : 107, as Tethys. 
Pruvot-Fol, 1934 : 39' as tigrinella Gray, 1850. 
Sowerby, 1869 : pi. 6. 

Aplysia (Varria) morio Verrill, 1901 : 25 

Text-figs. 28 and 29 

Synonym : modesta Thiele, 1910 : 124, probably a juvenile. 

Distribution. First recorded from Bermuda, but occurs from Rhode Island to 
Florida and Texas on the mainland. 

Material examined. Numerous specimens from Bermuda and the eastern 
seaboard of the United States. 

This is a very large but not bulky species. Verrill records a living specimen 400 
mm. long and 145 mm. high. A specimen in the Naturhistoriska Riksmuseum, 
Stockholm (Text-fig. 28) measured 250 mm. long, 65 mm. wide and no mm. high, 
the great height being due to the upward extent of the enormous parapodia, and not 
to bodily size. These parapodia can be spread widely also, giving a breadth in the 



REVISION OF THE WORLD SPECIES OF APLYSIA 



329 




Fig. 28. Aplysia morio. Dorsal view to show the wide cephalic tentacles, short neck, 
enormous parapodia, small closed mantle, large, leaf-like anal siphon, ctenidium and 
apertures of the opaline gland. From a specimen in the Swedish National Museum, 
Stockholm, from Bermuda. x #. 



zool. 5, 10. 



17 



330 REVISION OF THE WORLD SPECIES OF APLYSIA 

second specimen of 186 mm. The animal is a good swimmer. Dr. R. B. Hill of 
Bermuda Biological Station writes that he saw it swimming on the surface of the 
sea. Colour deep umber brown or nearly black, without spots, but with some dark 
lines or stripes on the head and sides. The foot is dark, the mantle and inner surface 
of the parapodia very dark, the ctenidium sometimes striped dark brown and light 
alternately. All parts of the body, including the foot, are soft. 

Head and neck relatively short, especially the latter, but they appear wide owing 
to the lateral extension of the hemispherical cephalic tentacles, which are rolled on 
their edges and are continued to the sides of the mouth slit. The eyes are small, 
on light areas of skin, just antero-lateral to the bases of the rhinophores, which are 
short, rather stout, slit deeply, pointed at the apex and set very close together. 

Foot elongated and very narrow, rounded in front, prolonged posteriorly as a 
short elliptical tail. It is wrinkled and soft, not tough, the edges wavy and well 
defined. The penis sheath (Text-fig. 29c) is anchored by numerous strands of muscle 
laterally ; its bulbous distal portion is not greatly enlarged, and is dark brown, 
with a smooth lining. The penis (Text-fig. 29^) is short, thick, tapering, curled 
round at the apex, unpigmented. The enormous parapodia are high and rounded. 
They commence close behind the rhinophores in the 250 mm. specimen, only 13 mm. 
behind them and 20 mm. apart, 52 mm. in front of the mantle. The flaps are thin, 
wide and deep brown inside, without markings, fluted on the edges and joined 
posteriorly low on the foot where they are tied to the tail by a little keel. They are 
thus free, flexible and obviously natatory. They expose the whole of the dorsal 
area, but Verrill remarks that they can overlap in repose. 

Mantle relatively small, less than a third the total length of the animal. In young 
specimens there is a small mantle foramen, but this closes later and in the adult 
there is no sign of the closure. The anal siphon is large, thin, tubular and leaf -like. 
Specimens from Florida exhibited a purple colour in the mantle glands, so that 
purple is secreted by them. 

Mantle cavity widely open, ctenidium large and exposed. Opaline gland large, 
simple, multiporous. Genital aperture hardly covered by the mantle, pigmented, 
with a frilled edge. The genital groove has a dark overhanging lip, but is itself 
light. 

The shell cavity, as above mentioned, is closed. The shell (Text-fig. 29, a, b) is 
long and narrow, without a spire, with a shallow anal sinus, the anterior border 
not narrowed. The dorsal edge of the apex and sinus are recurved, and one specimen 
has a hooked apex. The surface in older specimens is ridged concentrically. 

Jaws light brown in colour, rounded at the ends and composed of tall curved 
rods. 

Radula (Text-fig. 29, e, f) large and also light brown. There are about 86 rows, 
with a formula 56 . 1 . 56. Rhachidian tooth short and broad, the head narrow and 
excavated, the sides of the plate shouldered, the posterior border slightly concave. 
The cusp is equal to or longer than the plate, finely denticulate, with somewhat 
larger basal denticles. The first lateral has the usual broad head and an almost 
triangular plate. The denticulations are numerous and fine, the basal denticles 
larger laterally and secondarily denticulate in some. Along the row the cusp lengthens 



REVISION OF THE WORLD SPECIES OF APLYS1A 



33i 




+ 34 



Fig. 29. Aplysia morio. 

(a) Shell in dorsal view. The calcareous matter had been dissolved out in the specimen, 
leaving only a gelatinous mass under the horny portion. There are strongly marked 
ridges. x f . 

(b) Dorsal and ventral views of the shell of a younger specimen from Florida, now in the 
American National Museum, Washington, U.S.A. The concentric ridges are not 
figured, x f . 

(c) Penis sheath with its numerous anchoring muscles. x f . 

(d) Penis sheath opened to expose the penis and sperm groove. x I. 

(e) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth, x 80. 
(/) Thirty-fourth lateral tooth. x 132. 



332 REVISION OF THE WORLD SPECIES OF APLYSIA 

and narrows, the denticulations increase and involve the large lateral denticles, 
more being added on their sides. The three outermost teeth are vestigial. The 
feature of this radula is the extreme denticulation, both primary and secondary. 

Caecum large and corrugated, appearing on the surface for 30 mm., lying flat, 
but inflated at the end. Digestive gland relatively small. 

Cerebral ganglia fused to form a band. Visceral ganglia joined. 

Specific Characters 

Large size, deep brown colour, soft skin, wide hemispherical tentacles, narrow 
foot with short tail, very large, free, thin, natatory parapodia, arising close behind 
the rhinophores, then becoming wide, rounded and fluted on the edges, to unite 
low down on the tail. Mantle and visceral regions small. No shell foramen in the 
adult. Large leaf-like anal siphon, large ctenidium, simple multiporous opaline 
gland. Shell long and narrow, without spire, strongly ridged, with hardly any anal 
sinus. Radula with numerous rows and more than 50 teeth on each side in a full 
row. All the teeth, except the few outermost vestigial ones, with elaborate denticu- 
lation. Penis sheath anchored by numerous muscle strands, plain internally ; 
penis short, stout, unpigmented. 

Western North Atlantic. 



Aplysia (Varria) oculifera Adams & Reeve, 1850 : 64 
Text-fig. 30 

Synonymy : dactylomela (pars) ; Macnae, 1955 : 226. 
lineolata Adams & Reeve, 1850 : 64. 
nodifera Adams & Reeve, 1850 : 64. 

Telhys allochroa ; O'Donoghue, 1929 : 19 (non Bergh, 1905). 
Tethys hirasei Baba, 1936 : 10. 

Distribution. Red Sea, Seychelles, Persian Gulf, Pakistan, Ceylon, Mauritius, 
South Africa, East Indies, Formosa, Riu Kiu Islands, Japan, Western Australia, 
therefore widespread in the Indian and North Pacific Oceans. 

Material examined. Specimens from the Red Sea, Persian Gulf, Pakistan, 
East Indies and Western Australia. 

Baba (1949 : pi. 4) gave a coloured figure of a Japanese specimen as hirasei. In 
1956 (p. 216) he described the egg strings as yellowish green, with two to three eggs 
in each capsule. 

Aplysias of medium size, but rather coarse and bulky. Baba (1936) records a 
specimen 130 mm. long when alive ; preserved, a full grown specimen is about 90 
mm. long, 27 mm. wide and 42 mm. high. Slender when young, with a long pointed 
tail, but the mature specimens are never as large as specimens of A. dactylomela 



REVISION OF THE WORLD SPECIES OF APLYSIA 



333 



with which the species has been confused. Colour pale olive green, dull green, 
yellowish brown or rich chocolate brown, without radiating lines, but covered all 
over, except on the mantle and foot, with small black or brown rings, subequal 
in size and averaging i mm. in diameter. The outlines of the rings are neat, occasion- 
ally star-like, but not merging into the background as in A. dactylomela, nor are they 
linked by anastomoses. There are numerous rings on the head and even on the 
tentacles, but no large ring in front of the rhinophores. The centre of a ring may be 
buff, greenish white or white, and chalk granules may occur in groups between the 





#&&5L 



W 



Fig. 30. Aplysia oculifera. 

(a) Shell in dorsal and ventral views. The calcareous layer has been indicated by a 
dotted line, x f . 

(b) Penis extruded, with the flap at its base. X 2. 

(c) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth, x 80. The 
1 6th lateral enlarged, x 132. 



rings. There may also be dark mottled blotches between the rings. The mantle is 
speckled or sooty but not rayed. The inner sides of the parapodia are paler, with 
vertical bars of brownish green alternating with clear areas. The foot is pale. The 
skin is soft, but firm, and tends to be rough on the mantle. 

Head and neck elongated, but strongly retractile. Cephalic tentacles large, broad 
and mobile, short and rolled when contracted. Rhinophores acutely pointed, slender, 
slit half-way down, strongly contractile and sometimes branched. 

Foot moderately broad, rounded anteriorly, rather tough and wrinkled, unpig- 
mented, sharply marked off from the sides of the body, with a pointed tail. Penis 
(Text-fig. 306) short and broad, but can elongate ; spatulate to pointed at the tip, 



334 REVISION OF THE WORLD SPECIES OF APLYSIA 

not spirally twisted, with a flap at the base which is characteristic of the species. 
Parapodia long, not very high, widely spaced anteriorly, thick, frilled on the edges, 
coming close together low down on the foot posteriorly, and either free or joined 
low down, forming a shallow platform. Dorsal slit wide open. 

Mantle rather large and tough. Occasionally a papilla is present, or a very small, 
nearly closed aperture (Baba, 1936). Anal siphon short, broad, tubular, frilled. 
Purple glands well developed. 

Mantle cavity of moderate size. Ctenidium large, without pigment. Opaline 
gland simple, large, with numerous apertures, but as in other species a group of 
cells may unite anteriorly to secrete by a common aperture larger than the others. 
Internally, fibrous bands bind down the gland cells and these have to be dissected 
off to expose the gland fully. Genital aperture large, with smooth border in the 
young, but thickened and corrugated in the adult. Genital groove deep. 

Shell (Text-fig. 30a) varying from narrow to broad, neat, ovate, deeply concave. 
A firm calcareous lining does not quite reach to the edge of the horny portion. Dorsal 
edge recurved. Apex small and pointed, but strongly calcified. Anal sinus shallow. 

Jaws rectangular, short and wide. Rods of moderate length, loosely packed, tips 
square or oblique, but not broadened. 

Buccal mass rather small for the size of the animal. Radula with about 60 rows, 
38.1.38 (Text-fig. 30c). Rhachidian tooth with wide basal plate, which in young 
specimens is deeply excavated posteriorly so that it appears to be resting on two 
diverging legs, but in mature animals it becomes straight. Head narrow, cusp 
longer than the base but narrow, with a few fine denticles and two pairs of larger 
basal ones. In older teeth the edges are worn almost smooth. First lateral broad, 
with short broad basal plate and the usual pattern of short denticles. Cusps lengthen- 
ing and narrowing to about the 17th tooth, then shortening. All the laterals denticu- 
late, the three to four outermost teeth slender and degenerate. 

Caecum small, just breaking surface on the digestive gland and lying flat. 

Cerebral ganglia fused. Visceral ganglia distinct in young specimens, fused in 
the adult, with large granular nerve cells. 

Specific Characters 

Medium size ; green or brown colour, with numerous small, discrete, regular 
rings on the head and body ; foot strong with pointed tail ; penis short and broad, 
with a lobe at the base ; parapodia free ; opaline gland partly simple, partly com- 
pound, multiporous ; shell with firm calcareous lining ; closed mantle papilla ; 
purple glands ; radula with denticulations on the long narrow cusps of the laterals. 

Indian Ocean and North-west Pacific. 

References {see also synonyms) 

Baba, 1949 : 25, pi. 4, as hirasei. 

Baba, Hamatani & Hisai, 1956 : 216, as hirasei (breeding habits). 

Eales, 1944 : 4, juvenile specimen. 

Macnae, 1957 : 2 9°- 

Pilsbry, 1895 : IIQ an d io 9 {nodifera), both as Tethys. 



REVISION OF THE WORLD SPECIES OF APLYSTA 335 

Aplysia (Varria) pulmonica Gould, 1852 : 223 

Text-fig. 31 

Synonymy : Tethys pulmonica Pilsbry, 1895 : 96. 
T.p. var. tryoniana Pilsbry, 1. c. 

Distribution. South-east Asia, islands in the South Pacific (Samoa, Hawaii, etc) 
and Australia. 

Material examined. Specimens from Cochin China, Hawaii and other Pacific 
Islands. 

The species was named by Gould from specimens collected at Samoa by the United 
States Exploring Expedition in the Pacific 1838-42. Pilsbry (1895) copied Gould's 
figure (pi. 18) as Tethys and added a variety tryoniana. 

Aplysias of moderate size, up to 150 mm. long when alive and about 85 mm. long, 
35 mm. wide and 49 mm. high when preserved. Rather broad and flat, almost 
oblong in shape. Skin moderately soft, not flabby, but may become nodular and 
like crocodile-skin in contraction, and may have calcareous granules embedded in 
it. Much black pigment is present, forming a reticulate pattern or lines, but the 
general colour is a dark bronze green, which dissolves out on preservation, leaving 
only the black pigment. Foot dark and may be speckled. Inner sides of the para- 
podia with dark and clear blotches, especially near the edges (Text-fig. 31a). On 
the back, between the anterior ends of the parapodia, the pigment may be arranged 
in longitudinal lines or bands. Mantle blotched all over with black, including the 
anal spout, sometimes with light round spots about 2 mm. in diameter, filled in 
with black between the spots. Mantle cavity floor and ctenidium lightly pigmented 
or pale. A specimen from Honolulu, collected by Dr. T. Mortensen with a normal 
specimen, was intensely black ; it is now in the University Museum in Copenhagen. 

Head broad and flat, neck short and thick. Cephalic tentacles short, rolled, 
sometimes separated dorsally by a median groove, giving the head a bilobed appear- 
ance, with large mouth flaps. Eyes small. Rhinophores close to the cephalic tentacles, 
short and stout, close together, strongly contractile, resembling those of dactylomela. 
Gould described them as " faintly ringed ", but this was probably due to contraction. 

Foot of moderate breadth, with rounded anterior edges, defined border and distinct 
pointed tail, tough and hard in contraction. Penis sheath small and not very muscular, 
with two small retractor muscles. Penis (Text-fig. 316) broad, flat, curved and 
acuminate, the sperm groove reaching its tip. Parapodia widely spaced anteriorly, 
of moderate size, commencing close behind the rhinophores, joined low down poster- 
iorly forming a small platform, or sometimes a low wall shutting in the mantle 
cavity. The dorsal area is fully exposed. 

Mantle of moderate size, with a small tubular foramen on a ringed papilla, with 
or without starry rays. Anal siphon small, tubular, frilled. Purple glands present. 



336 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Mantle cavity not large. Opaline gland compound, with a large aperture, sur- 
rounded by numerous small openings. Genital aperture smooth edged, genital duct 
not raised on the floor of the mantle cavity, genital groove well marked. 

Shell (Text-fig. 31c) oval, thick and flat. Horny and calcareous layers present, 
the deep yellow horny portion with strongly marked concentric lines. Apex oblique, 




Fig. 31. Aplysia pulmonica. 

(a) Inner side of parapodium to show light and dark areas and clear edges, (cf . dactylo- 

mela, which has branched vertical bands, Frontispiece.) 
{b) Penis in its sheath and partly everted, x 2 J. 

(c) Shell in dorsal and ventral views. Note the almost pointed anterior border, x f . 

(d) Jaw plate (cf. dactylomela, Fig. 7/). 

(e) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth. X 80. 



without spire, but turned downwards, together with the ventral border of the short, 
deep anal sinus. 

Jaws (Text-fig. 31^) rounded with short, rather broad rods, some serrated on 
their free edges. 

Radula (Text-fig. 310) with about 60 rows and a formula up to 50 . 1 . 50. Rhachidian 
tooth with short but broad basal plate, slightly excavated posteriorly and deeply 
so anteriorly. Cusp longer than the plate in young teeth, with fine denticles which 
wear down in older teeth, and a rather large basal denticle on each side, which may 



REVISION OF THE WORLD SPECIES OF APLYSIA 337 

become subdivided. First lateral with short squat plate, finely denticulate broad 
cusp and the usual large basal denticle laterally. Remaining laterals with straight 
cusps of increasing length up to the middle of the row, almost smooth, with stout 
blunt basal denticle. Outermost three to five teeth vestigial. 

Caecum inflated at the tip, recurved or straight. 

Cerebral ganglia band-like, visceral ganglia joined but not completely fused. 

Specific Characters 

Moderate size, low broad shape, reticulate skin pattern, broad or moderately 
broad foot with pointed tail, broad flat penis, small parapodia joined low down, 
tubular foramen with starry rays in mantle, compound opaline gland with large 
aperture surrounded by small separate openings, broad oval shell with oblique apex, 
simple radula, the outer laterals almost smooth and spiky. 

Western Pacific from Asia to Australia. 

Note. Some specimens in contraction have a resemblance to A. dactylomela, 
but are lower and flatter, without rings on the skin, with a circlet of apertures to 
the opaline gland in addition to the main aperture, with a differently shaped penis, 
jaw plate, etc. Pulmonica and dactylomela occur in the same region in Honolulu 
and can be readily distinguished. Farran (1905 : 349) suggested that pulmonica 
and cornigera might be the same species. For differences between the two species 
see p. 303. 



Aplysia (Varria) rehderi sp. nov. 
Text-figs. 32 and 33 
Distribution. Monterey, California, U.S.A. 

Material examined. A single specimen in the U.S. National Museum (The 
Smithsonian Institution), Washington, U.S.A. The species is named in honour of 
Dr. Harald Rehder, Curator of Mollusks, U.S. National Museum. Holotype U.S.N.M. 
575055. 

The specimen measures 116 mm. long, 40 mm. wide and 62 mm. high, and is 
therefore of moderate size. There is little trace of colour, mostly black pigment on 
the inner sides of the parapodia. Skin thick and rather tough, probably much 
contracted. The general shape is low and rounded (Text-fig. 32, a, b). 

Head rather large and low, the neck narrower than the head. Cephalic tentacles 
(Text-fig. 32c) short and ear-like, free from the mouth slit. Eyes minute. Rhino- 
phores slender, tapering, fairly close together. 

Foot narrow, thick, smooth, 28 mm. broad at the broadest part, rounded in front, 
prolonged into a well developed pointed tail, projecting 12 mm. behind. Penis 
(Text-fig. 32^) enormous, thick at the base, at least 36 mm. long, spirally twisted, 
narrowing at the apex, the groove extending to its tip. Genital groove deep on the 



338 



REVISION OF THE WORLD SPECIES OF APLYSIA 



penis itself and on the muscular part of the stout, thick- walled sheath, the distal 
(bulbous) part of which is relatively small and smooth-lined. Parapodia small, 
tight, arising 16 mm. apart, not very high, joined behind, forming a low wall about 
8 mm. high around the mantle cavity. They are probably much contracted. 

Mantle long and narrow, 57 x 22 mm., thin, with a papilla at the point of closure, 
a good overhang anteriorly, but not much over the ctenidium. Anal spout short, 
broad, with sinuous margin. It is not possible to determine the colour of the secretion 
of the mantle glands. 




(a) 

(*>) 
(c) 
(d) 

(e) 



Fig. 32. Aplysia rehderi. 
Lateral view, x f . 

Dorsal view. Note the narrowed neck region, x f . 
Head in ventral view. 
Penis and penis sheath, x 1. 
Shell in dorsal and ventral views, x #. 



REVISION OF THE WORLD SPECIES OF APLYSIA 339 

Mantle cavity not large. Ctenidium very large and thick, 34 x 16 mm., curved 
round into the anal siphon. Opaline gland of medium size, compound, uniporous. 
Genital aperture very large and swollen, smooth, hardly covered by the mantle, 
genital groove deep. 

Shell (Text-fig. 320) 46 x 33 mm., broadly ovate, shallow, of unusual shape owing 
to the extreme narrowing at the apex. Horny and calcareous parts present, the former 
delicate with concentric ridges. No spire. Anal sinus long and deep. 

Jaws large and rectangular. Rods long, packed tightly together, rounded at the tips. 

Buccal mass very large, the radula (Text-fig. 33) being the largest of any species 
examined. It was 15 mm. long and 15 mm. broad, with 70 rows and a formula 
56. 1 . 56. Rhachidian tooth with a short broad basal plate, rather shapeless and not 
incised, the cusp broad, shorter than the base, with a few indistinct denticles and 




Fig. 33. Aplysia rehderi. Radular teeth. Rhachidian, first, sixth, 18th 
and outermost lateral teeth, x 80. 



broad denticulate basal denticle. First lateral short and broad, with curved basal 
plate and broad cusp shorter than the plate, its denticles small and fine. The cusps 
get narrower and longer along the row, and are almost straight down from the head, 
which is swollen and high. Denticles fine near the tip, mesially a few irregular ones 
and laterally large curved prongs, three to five in number, or even more, the most 
distal nearest the tip the longest, but very variable and irregular. The outer laterals 
with slender narrow plates and shorter cusps. Up to three vestigial teeth at the end 
of the row. 

Cerebral ganglia fused to form a band. 

Specific Characters 

Thick skinned, with swollen head owing to the large size of the buccal mass and the 
penis. Foot narrow, shorter anteriorly than the head, prolonged posteriorly into 
a pointed tail. Parapodia thick, joined posteriorly to form a low wall around the 
mantle cavity. Mantle thin, with conical papilla. Ctenidium large ; opaline gland 
compound, uniporous ; genital aperture and duct exceptionally large. Shell broadly 
ovate, shallow ; apex narrow, without spire or " accessory plate "; anal sinus 
long and deep. Radula 70 x 56 . 1 . 56, very large and broad, simple in the rhachidian 
and first few lateral teeth, then becoming elaborate in denticulation. Penis very 



34© REVISION OF THE WORLD SPECIES OF APLYSIA 

large and stout, spirally twisted, narrowed at the apex ; penis sheath smooth-lined 
distally. 
California. 

Aplysia (Varria) reticulata sp. no v. 

Text-fig. 34 

Distribution. North-west Australia, Queensland, Dampier Archipelago. 

Material examined. Five specimens from these areas, three from Queensland 
in the Australian Museum at Sydney, two in the British Museum. Type B.M. (N.H.), 
86.2.22.29 from N.W. Australia. 

Rather small Aplysias, the larger of the British Museum specimens measuring 
44 mm. long, 25 mm. broad and 29 mm. high when contracted. Skin firm and 
wrinkled, dirty yellow, with solid black spots scattered on the sides, and branched 
and anastomosing bands of dark pigment on the inner faces of the parapodia. Mantle 
roof in all the specimens with small black spots linked by less dense black anasto- 
mosing lines forming a network (Text-fig. 34, a-d). 

Head and neck small but contracted. Cephalic tentacles continued to the sides 
of the mouth slit. Rhinophores close together, stout, not as close together as in 
A. Sydney ensis. 

Foot narrow, only 10 mm. wide, very muscular and tough, with a short pointed 
tail. Penis (Text-fig. 342) short, flat, moderately broad, tapering. Parapodia large, 
arising close together far forwards on the neck and rising high above the mantle, 
then sloping towards the tail, but hardly joining posteriorly low down. The edges 
are fimbriated and thin. 

Mantle rather small and thick, 27 x 15 mm. There is a minute aperture in one 
specimen, but in another the shell sac is closed. Anal siphon of moderate size, 
frilled, projecting backwards between the parapodial lobes. Purple glands are 
present. 

Mantle cavity large and open. Opaline gland simple, multiporous. Genital 
aperture not enlarged, crescentic. 

Shell (Text-fig. 34/) 24 x 17 mm., delicate, with broken calcareous lining, elongated 
oval in shape, without distinct spire, deeply concave ; anal sinus wide but not 
deep. 

Jaws with very long, light brown, curved rods, closely packed together. 

Buccal mass small. Radula (Text-fig. 34^) 8x5 mm., rather narrow ; 50 rows, 
25 . 1 . 25. Rhachidian tooth with broad basal plate, slightly excavated on the poster- 
ior border and deeply so anteriorly. Cusp broad and blunt, almost as long as the 
base, with weak denticles. Laterals with short, rather square bases and more deeply 
cut denticles. One mesial denticle and two lateral denticles are larger than the others. 
Outermost two or three teeth degenerate. A common type of radula. 

Caecum curved and inflated at the tip. 

Cerebral ganglia fused, visceral ganglia joined. 



REVISION OF THE WORLD SPECIES OF APLYSIA 341 

Specific Characters 

Small size, spotted skin, reticulate pattern on the mantle roof, narrow foot with 
short tail, short flat penis, high free parapodia, minute mantle aperture or none, 
purple glands, simple multiporous opaline gland, deeply concave delicate shell, 
chief ganglia joined. 

Australia. 



oO a§ 

OOOO o 
• 




Fig. 34. Aplysia reticulata. 

(a) Pattern on the inner side of the parapodium. Edge light, then vertical alternating 
bands of light and dark, with light blotches. 

(b) Pattern on the mantle roof. Reticulate black lines with enlargements at the junc- 
tions. 

(c) Pattern on the side of the body. Oval or round black spots, interspersed with more 
numerous light areas on a dull yellow ground. 

(d) Specimen from the Dampier Archipelago, Australia, in lateral view, x § . 

(e) Penis sheath and its retractors, and sheath opened and extended to show the penis. 
X 2. 

(/) Shell in ventral and dorsal views, x § . 

(g) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth, x 80. 



342 REVISION OF THE WORLD SPECIES OF APLYSIA 

Aplysia (Varria) robertsi Pilsbry, 1895 : 89, pi. 55 (as Tethys) 

Distribution. West coast of North America, from Mexico to Central America. 

Material examined. One specimen from the west coast of Central America, 
now in the British Museum (Natural History). 

Of moderate size, about no mm. preserved, the B.M. (N.H.) specimen being 
93 mm. long, 35 mm. wide and 44 mm. high. Slender, soft and flabby. Pilsbry 
describes the colour as dirty light olive, with black-brown reticulations on both 
sides of the parapodia. Foot sole black. Mantle clear olive, the free border and 
siphon black. The specimen examined had the remains of black vertical stripes 
on the inner faces of the parapodia, and there was black pigment on the mantle 
roof and ctenidium, but not on the mantle floor. 

Head and neck elongated, the neck wider than the narrow head. Cephalic tentacles 
very small, slightly rolled on the edges. Rhinophores contracted, but thick, close 
together and set far back. Eyes plainly visible, antero-lateral to the rhinophores. 

Foot wide accordingly to Pilsbry, but strongly contractile and may appear narrow. 
Soft and fleshy, not strongly muscular. Tail long and slender. Penis sheath small, 
with a single retractor muscle, smooth-lined in the portion near the attachment of 
the small, flat, pointed penis. Parapodia rounded, thin, sinuous on the margins, 
arising far back, spreading widely, and meeting, but not joining low down posteriorly 
over the foot, forming an extension of the mantle cavity floor, which is not quite 
flat. 

Mantle 38 X 24 mm. in the 93 mm. specimen, with a minute papilla at its point 
of closure over the shell. Anal siphon broad and high. Purple glands present. 

Mantle cavity exposed. Opaline gland small, compound, with a single crescent- 
shaped aperture. Genital aperture not inflated, smooth edged. 

Shell delicate, thin and fragile, oval, almost flat ; 35 x 28 mm. Apex weak, with 
narrow, reflexed margin, anal sinus long and shallow (Pilsbry, 1895 : pi. 55, fig. 5). 

Radula 7x5 mm., flat, with 47 rows and formula 28.1.28. Narrower and less 
pointed than that of brasiliana. Rhachidian tooth with moderate, rather high basal 
plate, straight posterior border, short cusp a little longer than the plate, with well 
marked denticles, the basal denticle pronounced as usual. First lateral with short 
straight plate, short broad cusp and sharp denticles. Other laterals with longer 
cusps and elaboration of the denticles, all of which are deeply cut. Three to four 
vestigial teeth at each end of the row. 

Caecum small and appearing flat on the surface of the digestive gland. 

Cerebral ganglia fused, visceral distinct. 

Specific Characters 

Moderate size, slender flabby body, short cephalic tentacles, long tail, fleshy foot, 
freely mobile, thin parapodia, minute mantle papilla, small compound opaline 



REVISION OF THE WORLD SPECIES OF APLYSIA 343 

gland with one aperture, oval delicate shell, strongly denticulate radula, short flat 
pointed penis. 

West coast of North and Central America. 

The species bears considerable resemblance to A. brasiliana, but differs in the 
small size of the cephalic tentacles and the opaline gland, the shape of the genital 
duct beneath the mantle floor, the narrower radula with its more deeply cut denticu- 
lations, and the shape of the short, flat pointed penis. Pilsbry's specific name has 
therefore been retained until more information and specimens are available. 



Aplysia (Varria) sagamiana Baba, 1949 : 26, pi. 4 

Text-fig. 35 
Distribution. Japan. 

Material examined. Seven specimens from the Naturhistoriska Riksmuseum, 
Stockholm, collected by Dr. T. Gislen's Pacific Expedition, 1930-31, at Misaki, 
Sagami Bay, in 2-3 fm., on Zoster a. 

This pretty little Aplysia was first taken by His Imperial Majesty the Emperor 
of Japan and named and figured in colour by Baba, who also sketched the shell and 
radula. 

The largest Swedish specimen measures 31 mm. long, 10 mm. wide and 16 mm. 
high. Baba gives 70 mm. as a typical length, so that all specimens are small. Alive 
they are pinkish red in colour, with black spots on the head, neck, sides and parapodia. 
These spots are small, and may have the appearance of interrupted black margins 
to the parts concerned, distinguishing the species from A. parvula, in which the 
margins are continuous. The cephalic tentacles and rhinophores are tipped with 
black, the mantle is mottled. The general shape is slender, high and narrow, the 
skin soft. 

Head small, neck long. Cephalic tentacles wide, slender, rolled on the edges, 
fluted in some specimens, continued to the ventral sides of the mouth slit. Eyes 
plain but small, close to the base of the rhinophores, which are small, slender, acutely 
pointed and slit as usual. 

Foot narrow, only 6 mm. wide in a 27 mm. specimen, soft, rounded in front, a 
little wider at the middle, with narrow pointed tail and the edges plainly marked 
against the pigmentation of the sides of the body, for the foot is colourless. Penis 
(Text-fig. 356) of the short, broad type, in a smooth-lined and not enlarged sheath. 
Parapodia well developed, rounded, with fluted margins, the inner sides with a block 
pattern of dark and light areas (Text-fig. 35a), capable of closing over the dorsal 
region or of spreading wide open, joined low down on the foot posteriorly, forming 
a flat continuation of the mantle cavity floor. 

Mantle 12 x 8 mm., soft and thin, with a small flat aperture, rayed. Siphon 
rounded and low. Purple secretion exuded. 

Mantle cavity of moderate size, open at both ends. Ctenidium small and neat, 
opaline gland small, simple, multiporous, but there is a tendency for the anterior 



344 



REVISION OF THE WORLD SPECIES OF APLYSIA 



vesicles to be grouped and for their ducts to join to form a larger aperture than the 
others. Genital aperture under cover of the mantle rim, genital groove small. 

Shell elongated, oval, 10 x 7 X 2 mm., not strongly concave, delicate, without 
spire, apex acute, oblique, edge slightly recurved ; anal sinus well formed. 

Jaws composed of flat-topped rods, often with a little projection at one corner. 

Radula (Text-fig. 35^) 29 x 18.1.18, but Baba gives 35 x 22.1.22. Rhachidian 
tooth with short but broad basal plate, the anterior border unusually broad and 
deeply incised. Cusp as long as or slightly longer than the base, with about seven 
neat graduated denticles on each side, the basal ones being large. In older teeth 






X&&V 



Fig. 35. Aplysia sagamiana. 

(a) Block pattern on inner side of a parapodium. 

(b) Penis in its sheath. X 10. 

(c) Caecum exposed on the surface of the digestive gland and lying flat. 

(d) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth, x 132. 



the fine denticles are worn down, but the basal denticles become accentuated. First 
lateral with a bulbous head and short tapering basal plate, the denticulations 
resembling those of the rhachidian, but the mesial basal denticle is smaller. Along 
the row the basal plates become elongated and curved, the head narrows, the cusp 
elongates and so does the penultimate lateral denticle. After the middle of the row 
the cusps shorten and eventually disappear, until the three to five outermost vestigial 
teeth are reached. This is a neat, well denticulated radula. 

Caecum lies flat on the surface and is somewhat inflated at the apex (Text-fig. 35c). 

All the nerve ganglia are distinct and separate, but the cerebral ganglia are 
contiguous, without a visible commissure. 

Specific Characters 

Small, with interrupted or speckled black markings on a pinkish red ground ; 
long cephalic tentacles, narrow foot and slender tail ; short broad penis ; relatively 



REVISION OF THE WORLD SPECIES OF APLYSIA 345 

large parapodia, fluted at the edges, mobile, joined low down on the foot posteriorly. 
Mantle thin, with a small flat rayed aperture. Purple secretion. Opaline gland 
simple, multiporous, but with a tendency to fusion of the anterior vesicles. Shell 
delicate, shallow, with acute oblique apex. Radula small, up to 35 rows and formula 
22 . 1 . 22, rhachidian tooth broad, deeply incised anteriorly, with clear cut and regular 
denticles, laterals also with neat graded denticles. All the nerve ganglia distinct. 

Japan. 

There is some resemblance between this species and A. parvula, but the pattern 
and markings, the low junction of the parapodia, the small mantle aperture, the 
shallow shell without spire and the broad rhachidian tooth of the radula distinguish 
it. One resemblance, however, is in the primitive separation of the chief nerve 
ganglia, so that this species forms a link between the subgenus Pntvotaplysia and 
the main group in which the cerebral ganglia are fused. 



Aplysia (Varria) sowerbyi Pilsbry, 1895 : 101 
Text-fig. 36 

Synonymy : A. tigrina (non Rang, 1828) ; Angas, 1867 : 228 ; Sowerby, 1869, pi. 2. 

Distribution. Australia, New South Wales. Victoria. 

Material examined. Four specimens from Long Reef, Sydney, New South 
Wales, October 1957, E. Pope and party. 

The four specimens are small, the largest being 67 mm. long, 16 mm. wide and 
27 mm. high, with parapodia spread 38 mm. wide, foot 12 mm. wide. They bear 
a general resemblance to sydneyensis in their slight build. Colour yellowish green 
with small dark brown blotches, or dark greenish brown with light, almost white, 
spots and faint brown markings. There is an all-over brownish black reticulation, 
either making a honeycomb pattern or drawn out near the edges of the parapodia 
into parallel vertical lines. The preserved specimens have neither colour nor markings 
on the foot, inner sides of the parapodia, mantle, or mantle cavity floor, except for 
a tinge of grey over the opaline gland. Skin soft and flabby. 

Head small, with short neck (Text-fig. 366). Cephalic tentacles small, curved 
backwards, almost sickle shaped, with small connections with the sides of the mouth ; 
the shape is characteristic. Eyes large and conspicuous. Rhinophores very long 
(12 mm.), set very close together, slit deeply. 

Foot narrow, soft, with rounded anterior edges, not well marked off laterally 
from the sides of the body, posteriorly with a short triangular tail about 5 mm. 
long. Penis sheath small, recurved; with two small retractor muscles (Text-fig. 
36^). Penis filiform, small (Text-fig. 36^). Parapodia not large, soft and thin, mobile, 
arising close to the rhinophores, rounded, with slightly sinuous edges. Posteriorly 
they do not unite or meet, but stand up like the edges of a rounded collar, with 
about 3 mm. space between them, the mantle cavity being therefore open posteriorly. 

ZOOL. 5, IO. l8 



346 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Mantle thin, narrow, pointed anteriorly, with a thin flap laterally where a purple 
gland lies, and a soft, broad, rounded anal spout, which in one specimen is expanded 




(a) 
(b) 
(o) 
(d) 

(') 



Fig. 36. Aplysia sowerbyi. 
Dorsal view of specimen with expanded parapodia. x 



Head of another specimen in anterior view. X 1 J. 

Tail in lateral view to show upstanding parapodia, which do not join or meet, x 

Penis sheath and its two weak retractor muscles, x ij. 

Penis, x 2. 
(/) Shell in ventral view, x 1 J. 
(g) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth, x 132. 



ij. 



and leaf -like. There is a minute papilla at the point of closure over the shell, probably 
open during life. 

Mantle cavity open at both ends. Genital aperture small, smooth, not inflated, 
inconspicuous. Opaline gland large, reniform, slightly pigmented, with one larger 
and numerous small apertures of the predominantly simple gland. Ctenidium small, 
unpigmented. 



REVISION OF THE WORLD SPECIES OF APLYSIA 347 

Shell (22 x 16 mm.) as figured by Sowerby and copied by Pilsbry (Text-fig. 36/). 
It is firm and horn coloured, with distinct growth rings in the form of ridges. Shape 
narrow and concave, with dense but smaller calcareous lining and a hard, strongly 
calcified and oblique, incurved apex. It is unlike the delicate shell of Sydney ensis 
or the almost flat shell of cronullae. 

Jaws 4x3 mm., curved, brown. Rods long, almost straight, rounder and a little 
wider at the tip. 

Radula 7x5 mm., with about 45 rows and formula 26.1.26 (Text-fig. 36^). 
Rhachidian tooth with broad basal plate, slightly excavated posteriorly and incised 
anteriorly but with a small projection centrally. Cusp about equal in length to the 
plate, strong, with well marked and rounded denticles, the two basal ones of each 
side being enlarged. First lateral with a narrow triangular basal plate, broad head 
and short cusp, the denticles repeating those of the rhachidian tooth. Along the 
row the basal plates elongate and become curved, but not very broad, the cusps 
lengthen and are narrow and spiky, longer than the plates. Their denticles are 
weak near the apex, but there is one strong mesial denticle and laterally there are 
at least three, of which that nearest the apex is the largest and may be nearly half 
the length of the cusp, giving a two-pronged appearance to the teeth. From three 
to four of the outermost teeth are small and vestigial. 

Caecum appearing flat on the surface of the digestive gland. 

Cerebral ganglia fused, flat, band-like ; visceral ganglia distinct. 

Specific Characters 

Medium size, slightly built. Ground colour yellowish green to dark brownish 
green, with small dark blotches, sometimes light spots and usually with dark reticu- 
lations. Pale on the foot, inner sides of the parapodia, mantle and lining of the 
mantle cavity. Skin soft. Head small, neck short. Cephalic tentacles short, sickle- 
shaped. Rhinophores long, deeply slit, set very close together. Foot narrow, with 
short tail. Penis small, filiform, in a small, recurved sheath. Parapodia thin, mobile, 
not joining or meeting posteriorly. Mantle thin, with minute papilla. Purple glands 
present. Mantle cavity open at both ends. Opaline gland large, reniform, simple, 
with one large and numerous small apertures. Shell firm, concave, with strongly 
oblique hard apex. Radula 45 x 26.1.26, characterized by long spiky cusps of the 
laterals and narrow basal plates. 

Australia, New South Wales. 

The species is related to Sydney 'ensis and cronullae. It resembles the former in 
its slight build, small head and cephalic tentacles, narrow foot, filiform penis, weak 
parapodia, simple multiporous opaline gland and radular formula. Its resemblances 
to cronullae are the filiform penis, large opaline gland, small mantle papilla and radular 
formula. It differs from both in the absence of pigment on foot, inner sides of the 
parapodia, mantle and mantle cavity floor, in the fusion of some of the opaline 
glands to discharge by a larger anterior aperture than the small ones of the isolated 
gland cells, in the hard oblique apex of the firm, concave shell and the long spiky 
lateral radular teeth. 



348 REVISION OF THE WORLD SPECIES OF APLYSIA 

Aplysia (Varria) Sydney ensis Sowerby, 1869 : pi. 7 
Text-fig. 37 

Synonym : A. excavata Sowerby, 1869 : pi. 3. 

Distribution. Australia : New South Wales, Victoria, South Australia, Western 
Australia, Queensland. 

Material examined. Specimens from all the above areas. 

This species was named by Sowerby from a shell, and has never been clearly 
defined. Australian zoologists recognize the species, and this account is based on 
specimens named by them in their museums. 

Aplysias of moderate size, usually about 125 mm. long, shrinking to 80 mm. long, 
22 mm. wide and 28 mm. high when preserved. Rather low in shape and not bulky 
(Text-fig. 37, a, b). Colour dark brown to almost black, spotted all over with cream, 
grey or dark brown spots, with a black lined pattern on the head and sides, reticulate 
in places. The mantle has a characteristic pattern of dark dots scattered on a light 
ground and forming rings at intervals, or radiating stripes (contrast A. reticulata 
p. 340). Foot dark. Inner sides of the parapodia with black bands arranged vertically 
and lighter spots near the edges. No pigment on the ctenidium or the mantle cavity 
floor. Skin soft, wrinkled. 

Head small, neck short and thick. Cephalic tentacles short, pointed, rolled on 
the edges, not projecting much. Eyes small. Rhinophores tall, tapering and slender, 
very close together. 

Foot narrow to moderate, contracting with irregular wrinkles, soft and flabby, 
anterior edges rounded, tail distinct, slender. Penis sheath small, penis short but 
narrow and pointed (Text-fig. 37c). Parapodia mobile, not large, commencing close 
to the rhinophores, rounded at first, thin, their edges sinuous but not fimbriated, 
meeting low down over the foot posteriorly, forming a platform-like extension of 
the mantle cavity floor. The dorsal area is fully exposed. 

Mantle rather small, long and narrow, thin, with an aperture about 1 mm. in 
diameter marked by black radiating lines, but in a much contracted specimen this 
may be reduced to a small papilla. Anal siphon tubular, rounded, or tall and leaf- 
like. Purple glands present. 

Mantle cavity widely open. Ctenidium of moderate size. Opaline gland small, 
pyriform, simple, multiporous. Genital aperture small, crescentic, covered by the 
mantle edge, genital groove delicate. 

Shell (Text-fig. 37^) thin, pale in colour, narrowly oval and concave, with horny 
and calcareous matter, and an anal sinus, which is very short, posteriorly placed and 
shallow. Apex narrow and hard, lines of growth pronounced. 

Jaws dark brown, almost triangular in shape. Rods short to moderate, curved at 
the tip, tapering. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



349 



Buccal mass small, radula acutely pointed, 7x7 mm. (Text-fig. yje). Formula 
about 45 X 28 . 1 . 28 but usually less. All the teeth except the outermost vestigial 
ones denticulate, with rather short basal plates and heavily built heads. Rhachidian 
tooth sunken and pale, with a short and moderately broad shouldered basal plate, 
excavated anteriorly and almost straight posteriorly, the cusp shorter than the 





Fig. 37. Aplysia Sydney ensis. 

X I. 



(a) Dorsal view of a specimen with expanded parapodia. 

(b) Lateral view of another specimen, x f . 

(c) Penis sheath opened to show the narrow penis. X if. 

(d) Shell in dorsal and ventral views. The fine concentric ridges are not shown, x § . 

(e) Radular teeth. Rhachidian, first, sixth, 12th and outermost lateral teeth, x 80. 



plate and bearing feeble denticles, of which the basal pair are usually larger than 
the others. The first lateral also has irregular denticles on a short cusp, a broad 
head and out-turned basal plate. Further along the row the cusps remain broad 
but do not increase greatly in size, although their plates become narrower. The 
mesial basal denticles tend to form a projecting ridge on the edge of the cusp and 
the lateral basal denticles are enlarged and may be secondarily denticulate. The 
outermost three teeth are vestigial. 

Caecum straight, just reaching the surface of the digestive gland near its tip. 

Cerebral ganglia fused, visceral still separately recognizable. 



350 REVISION OF THE WORLD SPECIES OF APLYSIA 

Specific Characters 

Moderate size, low shape, pattern of lines and spots, small head and short cephalic 
tentacles, narrow foot with slender tail, short narrow pointed penis, mobile parapodia 
fused low down on the tail, flat mantle aperture with radiating lines, simple opaline 
gland with many apertures, oval hyaline concave shell with shallow anal sinus, 
radula with few teeth in a row, short bases, enlarged heads and irregular denticles. 

Australia. 

The copepod Strongylopleura pruvoti Monod & Dolfuss, 1932 was found in the 
mantle cavity of three specimens from Sydney, now in the Naturhistorisches Museum, 
Wien. See also p. 368. 

References (see also synonym) 
Allan, 1932a : 419 ; 1950 : 213. 
Angas, 1877 : 190, as Sydney ensis and excavata. 
Hedley, 191 8 : 107, as Tethys sydney 'ensis and excavata. 
Hutton, 1882 : 118. 
Pilsbry, 1895 : 100, as T. excavata ; 101, as T. sydney ensis. 



Aplysia (Varria) willcoxi Heilprin, 1886 : 364 

Text-fig. 38 

Synonym : Tethys willcoxi var. perviridis Pilsbry, 1895 : 81. 

Distribution. East coast of North America, from New England to the West 
Indies. Recorded from Woods Hole, Mass.; New Jersey ; Rhode Island ; Dry 
Tortugas ; Florida and Guadeloupe. 

Material examined. Specimens from Woods Hole and Fisher Island, New 
York. 

First recorded by Heilprin from Florida, this species is now the most northerly 
one known on the East Atlantic coast of North America. It is a large, very bulky 
Aplysia, about 200 mm. long, but contracting considerably when preserved (Text-fig. 
38a). A specimen from Fisher Island measured no mm. long, 40 mm. wide and 
54 mm. high. Skin smooth but tough and leathery. Colour variable, greenish yellow 
ground colour with irregular purplish black marbling on the parapodia, neck and head, 
sometimes with round dark spots or blotches and light areas on the inner sides of 
the parapodia, but these are not regular. Foot, mantle and ctenidium dark. Plain 
deep seagreen specimens were described as var. perviridis. Green specimens may 
have a reticulate black pattern with some white spots. When preserved the markings 
are obliterated and much green pigment is exuded into the preserving fluid, the 
whole animal appearing deep green in colour. Sanford (1922) states that the animal 
has " a slightly unpleasant odour ". 



REVISION OF THE WORLD SPECIES OF APLYSIA 



35i 



Head and neck large. Cephalic tentacles well developed, broad, curved, rolled 
at the edges. Rhinophores set well back and fairly close together, conical, with short 
slits. 

Foot narrow, 30 mm. wide anteriorly, narrowing to 25 mm. Front edges rounded, 
sides ill defined, tail short. Penis very long, filiform, 45 mm. long in the no mm. 
specimen. (Pilsbry gives a figure of a penis only partly extruded from its sheath and 
describes the latter as the penis itself.) Parapodia large, thick, natatory, arising 
close to the rhinophores, borders smooth or slightly irregular but not fimbriated, 
joined low down over the foot. 




b) 



(a) Lateral view. 

(b) Radular teeth. 




Fig. 38. Aplysia willcoxi var. perviridis. 



Rhachidian, first sixth and 17th lateral teeth. X 80. 



Mantle large with minute tubular foramen, sometimes surrounded by wrinkles 
or short black radiating lines. Anal siphon long, wide and tongue-like, but contractile. 
Purple glands exude a reddish purple fluid which turns crimson in formalin. 

Mantle cavity with large ctenidium, compound uniporous opaline gland and large 
inflated genital aperture with smooth edge. 

Shell broad, flattened, thin, with some calcareous matter. It measured 56 X 43 
mm. Concentric and radiating ridges marked, apex small, curved, anal sinus long 
but shallow, with a thickened reflected margin. 

Jaws large, composed of closely packed, short, slightly curved rods. 

Radula (Text-fig. 386) large, 11 x 10 mm. About 60 rows and a formula of 
30.1.30. Rhachidian tooth feeble, lighter than the other teeth, the basal plate 
rather long, with straight anterior and posterior borders and short rounded cusp, 



352 REVISION OF THE WORLD SPECIES OF APLYSIA 

bearing weak denticles, the basal one, as usual, larger than the others. First lateral 
with short basal plate, large head and short cusp, upon which the median basal 
denticle is well developed, as well as two or three lateral ones, but the apical denticles 
are mere nodules. The basal plates of the remaining lateral teeth become squared 
off, but the cusps rarely exceed the plates in length. A striking feature of this radula 
is the elaboration of the basal denticles on each side of the cusp. Thus the 17th 
lateral has four well developed basal denticles mesially and three or four large ones 
laterally, although the denticles of the main cusp remain feeble. The three outermost 
teeth are vestigial. 

Caecum small, only its tip appearing on the surface of the digestive gland. 

Cerebral ganglia fused to form a flat band. Visceral ganglia joined but not fused. 

Specific Characters 

Large size, bulky shape, tough skin, dark colouring of all parts with abundance of 
deep green in the var. perviridis. Cephalic tentacles large, rhinophores close together, 
foot narrow, penis long, filiform, parapodia smooth, natatory, joined low down 
posteriorly, minute mantle foramen, purple glands, opaline gland compound uni- 
porous, radula large with elaboration of the basal denticles. 

North-west Atlantic. 



Merriman, 1937 : 95. 
Sanford, 1922 : 80. 
White, 1952 : 112. 



References {see also synonym) 



Aplysia (Varria) winneba Eales, 1957 : 180 
Distribution. West coast of Africa : Ghana, near Accra ; Cape Verde Islands. 

Material examined. Numerous specimens from this area. Colour photographs 
of the living animal swimming in a rock pool were lent by Mr. R. Bassindale, who 
collected many of the specimens. 

LectotypeB.M. (N.H.), 1957.5. 14. 1. (Eales, i%jProc.nialac. Soc. Lond. 32 : 208). 

Aplysias of medium size, highly mobile, a typical specimen, preserved, measuring 
70 mm. long, 55 mm. wide and 45 mm. high, being therefore broader than high 
when the parapodia are spread. When strongly contracted, however, the small head 
is drawn in, the parapodia are contracted tightly over the mantle but do not quite 
close the dorsal slit, and the inflated sides of the body make the animal appear like 
a little balloon. A large number of the specimens were in this condition. The general 
colour is purplish black, with browner shades on the mantle, vertical bars of black 
alternating with lighter areas on the inner edges of the parapodia, and a characteristic 
light triangular patch at the anterior end. When preserved the purple colour fades 
out, leaving a mottled pattern of grey and brownish black, with lighter areas between. 
There is no definite pattern and no rings are present. Some specimens have chalky 



REVISION OF THE WORLD SPECIES OF APLYSIA 353 

granules embedded in the skin, which is soft and firm. The foot and mantle floor 
are unpigmented ; the mantle may have radiating irregular lines or blotches of 
pigment. 

Head and neck small, but the neck is long. Cephalic tentacles short and broad 
when contracted, but expanded they stand out from the sides of the head, and have 
fimbriated edges. They are linked with the ventral sides of the mouth slit. Eyes 
small. Rhinophores slender and pointed, strongly contractile, slit half-way down. 

Foot of moderate width, about 20 mm. wide in a 90 mm. specimen. Tough and 
nodular in contraction, its borders defined and wavy. Tail short and tapering. 
Penis long and filiform, but can be inflated, penis sheath smooth lined. Parapodia 
not large, but strongly contractile. They arise by rounded edges rather far back, 
almost at the level of the genital aperture, and are spread wide in swimming, being 
flapped upwards alternately during this movement. They are joined low down 
over the tail, forming a kind of spout into which the anal siphon fits. Their edges 
are fimbriated. 

Mantle of moderate size (26 X 19 mm. in a 70 mm. specimen), thin, with a small 
open tube about J mm. in diameter. Anal siphon broad and frilled, with a tendency 
to form a spout towards the tail, or standing upright with reflexed edges. Purple 
glands present. 

Mantle cavity shallow. Ctenidium colourless, of medium size. Opaline gland 
compound, large, with conspicuous aperture. Genital duct raised on the mantle 
cavity floor, its aperture fimbriated. 

Shell 30 X 23 x 4 mm., of narrow oval shape, with remains of a calcareous layer. 
Apex without spire, but slightly recurved, and the dorsal side is turned back. Anal 
sinus rather long. 

Jaws of slender, slightly curved rods, tightly and neatly packed. 

Radula 8-5 x 7 mm., about 55 rows and a formula 37.1.37. The rhachidian 
tooth is high for its width, with a rounded cusp, irregularly denticulate. The first 
lateral has a similar rounded cusp, one larger denticle mesially and one laterally, 
in addition to those on the cusp itself. Along the row the cusp elongates and becomes 
narrower, the large mesial denticle persists and the lateral one enlarges, adding 
a small one nearer the base of the cusp. After the 16th tooth the cusps become 
simpler and smaller. The five outermost teeth are degenerate. This is a common 
type of radula in tropical Aplysias, but differs from most in the irregularity of the 
denticulations. 

Caecum inflated at the apex, lying straight on the surface of the digestive gland, 
with its tip bent inwards. 

Cerebral ganglia fused, visceral distinct. 



Specific Characters 

Aplysias of moderate size, purplish black in colour, with vertical bands of dark 
and light on the inner sides of the parapodial edges, and a triangular light patch 
anteriorly. Highly mobile and strongly contractile. Cephalic tentacles fimbriated, 
also the parapodia and anal siphon. Foot of moderate width with short tail. Penis 



354 



REVISION OF THE WORLD SPECIES OF APLYSIA 



long and filiform. Mantle thin with a small tubular aperture. Purple gland present. 
Opaline gland compound with a single conspicuous aperture. When preserved the 
animal may assume a short broad ballooned shape. 
West Africa. 



Subgenus Aplysia Linnaeus, 1767 

Aplysia (Aplysia) cedrosensis Bartsch & Rehder, 1939 : 2 

Text-figs. 39 and 40 

Distribution. Lower California. Named by the authors from Cedros Island in 
the Gulf of California. 

Material examined. The type, in the United States National Museum (No. 
472859), and another specimen, which had been in the same Museum unnamed since 
1889 and came from San Bartolome Bay, Mexico. 

This is one of the bulkiest members of the genus, and is so much enlarged posteriorly 
that it bears a superficial resemblance to a Dolabella. The type measures 165 mm. 
long, 90 mm. broad and 85 mm. high, but it must be much larger when alive. The 
general shape is low and very broad (Text-fig. 39a) . The skin is tough and leathery, 
rough and carunculate, black and brown all over, but described as grey originally, 
with black patches and an elongated black band dorsally posterior to the rhinophores. 
Mantle and inner sides of the parapodia mottled and blotched irregularly. The head 
is not much contracted, the neck thick, the parapodia short and exposing the whole 
of the dorsal area. 

The broad head is clumsy, the cephalic tentacles widely separated, hardly rolled 
at the edges, not continued to the mouth. Eyes small on clear areas of skin. Rhino- 
phores conical, stout, slit half-way down, wide apart. 

Foot very broad, 118 X 70 mm. in the type specimen, rounded in front, with a 
tendency to form lateral wings, the edges defined, the tail short, broad, round, 
strongly contracted and bent ventral wards, so that it is difficult to determine whether 
it was sucker-like ; it is thick and tough. The penis and penis sheath exhibit clearly 
the characters of the subgenus (Text-fig. 39, c, d, e). The penis is black, very large 
and stout, being 54 mm. long and 11 mm. broad at the base, but tapering to a point. 
The sperm groove extends nearly to its tip. The penis sheath is divided as usual 
into two portions, a ridged muscular proximal part near the aperture and a basal 
distal part which in this species, although swollen, is not large enough to contain 
the whole of the penis, even when it is bent back on itself. The bulbous portion 
is lined with warts which in the type specimen are simple and strong, in the second 
specimen more slender and sometimes branched. Each wart bears simple spines, 
slightly recurved and probably of a horny or chitinous composition. The parapodia 
are small, short, widely separated anteriorly, giving a broad neck region. They are 



REVISION OF THE WORLD SPECIES OF APLYSIA 



355 




Fig. 39. Aplysia cedrosensis. 

(a) The type specimen in dorsal view. Modified from the sketch by Bartsch & Render, 
I 939- The mantle is a little foreshortened and the mantle edge has been reflected. 
The ctenidium has been lifted out of the mantle cavity to show its size, x f . 

(b) Shell in ventral view. X f . 

(c) Penis sheath opened to show the bulbous warted region and the large penis, x •§. 

(d) Penis tip. X if . 

(e) Warts and spines from the distal portion of the penis sheath, x 21. 



356 REVISION OF THE WORLD SPECIES OF APLYSIA 

rather tight over the dorsal area and do not close it, and are very thick and rough. 
Posteriorly they unite to form a wall about 25 mm. high, shutting in the mantle 
cavity. 

Mantle short and broad, not covering either the ctenidium or the genital aperture. 
There is a small, round, flat foramen with radiating lines of contraction around it. 
The anal siphon, 20 x 35 mm., is broad but not high, strongly attached to the mantle, 
flat on the side of it and only free for 5 mm.; it is therefore non-tubular (cf. A. dura, 
p. 362). The edge of the siphon is irregular but not fimbriated. Glands are present 
under the small mantle flap, but no information is available regarding the colour 
of their secretion. 

Mantle cavity small, ctenidium large and projecting. Opaline gland of simple 
vesicles, multiporous. Genital aperture large and frilled, genital groove deep, with 
strongly developed dorsal lip. 




Fig. 40. Aplysia cedrosensis. Radular teeth. Rhachidian, first, sixth, 24th and 

35th laterals, x 80. 

Shell (Text-fig. 396) large and broad, 76 x 57 mm., thin and flat, dark brown in 
colour. Growth lines are well marked and there are radiating ridges from the apex. 
No calcareous matter is present, but Bartsch & Rehder report slight calcification. 
No spire, but a slight turn-over at the apex. Anal sinus short and shallow, with the 
broad outer angle characteristic of the subgenus. 

Radula (Text-fig. 40) very large, with 118 rows and a formula of 80 . 1 . 80. Rhach- 
idian tooth pale and sunken, rather high, with narrow head and broadening plate. 
Cusp shorter than the plate, with a pair of denticles near the tip ; these get worn 
away in older teeth, as does also the cusp itself. First lateral with a broad head and 
short plate, short cusp with one denticle on each side, not forming a pair, the mesial 
denticle nearer the point than the lateral one. This applies also to the other teeth in 
the row, all of which are simple, although they have longer and more curved plates, 
narrower heads and longer and more acute cusps. Some of the outer laterals may 
develop two denticles mesially, but on the whole the radula resembles closely that 
of A. Juliana. There are three vestigial teeth at the end of the row. 

Cerebral ganglia forming a fused band, visceral ganglia joined. 

Specific Characters 

Large bulky shape, much enlarged posteriorly, skin thick, rough, leathery, dark 
in colour and mottled and blotched with brownish black, but without regular pattern. 



REVISION OF THE WORLD SPECIES OF APLYSIA 357 

Head broad, flat and clumsy, foot broad, parapodia small, low, thick, joined posterior- 
ly to shut in the mantle cavity ; mantle foramen round, flat ; anal siphon non- 
tubular, wide, flat on the side of the mantle ; shell broad, flat, without spire, but 
with short shallow anal sinus and large outer angle ; radula large, simple, with 
many rows (more than 100) and numerous teeth in a row (up to 80) . No information 
on the secretion of the mantle glands, but the inference would be that they secrete a 
milky substance, not purple. 
Lower California. 



Aplysia (Aplysia) depilans Gmelin, 1791 : 3103 (see pp. 274 and 396). 

Text-fig. 41, a-e 

The type species of the genus. 

Synonymy : Laplysia depilans (= one kind of Lernaea of Bohadsch, 1761 : 49), 
and L. d. major Barbut, 1783 : 32. 
Dolabella fragilis Lamarck, 1822 : 42. 
Dolabella laevis de Blainville, 1819 : 395. 
Aplisia leporina delle Chiaje, 1822 : 28. 
A. poli delle Chiaje, 1822 : tav. 3. 
A. poliana delle Chiaje, 1822 : 30. 
Aplysia major Lankester, 1875 : 13. 
A. melanopus Couch, 1870 : 173. 
A. petersonii Gray, 1828 : 4, pi. 4 fig. 4 : Sowerby, 1869 : pi. 2. 

Distribution. Mediterranean, Atlantic coasts from France and the Channel 
Islands to West Africa (Morocco, Rio de Oro), Madeira. Rarely on the southern 
British coasts. 

Material examined. Specimens from the Mediterranean and French Atlantic 
coasts. 

The specific name is due to the erroneous belief that handling the living animal 
causes the hair to fall out. Rang (1828 : pis. 16 and 17) gave excellent figures of the 
animal. Mazzarelli's monograph on the Aplysiidae of the Gulf of Naples (1893) 
included an account of the three Mediterranean species, with coloured figures. In 
depilans (1891 : 3) he noted the spiral apex of the shell in the juvenile, the nodosities 
on the penis sheath and the spiral caecum. These nodosities had first been seen and 
recorded by Bohadsch (1761) and were figured by delle Chiaje (1828 : pi. 2). 

Large Aplysias, low and broad, in contrast to the narrowness and height of A. 
fasciata, bulky, attaining 300 mm. in length when alive, 200 mm. long, 100 mm. 
wide and 80 mm. high when preserved. Skin smooth and firm. Colour variable, 
brown to greenish brown, dark, with large white or grey blotches, and fine dark 



358 



REVISION OF THE WORLD SPECIES OF APLYSIA 



veining. There may be a violet border to the tentacles, rhinophores, parapodia 
and the foot sole. Mantle speckled brown. Inner sides of the parapodia blotched 
with dark and light patches. All colour may disappear with preservation. The animal 




b) 







Aplysia depilans. 



* 



Fig. 41. 

(a) Foot sole to show sucker, x f . 

(b) Penis sheath and retractor muscles, x 

(c) Penis sheath opened to show the penis and the nodosities on the lining of the distal 
portion of the sheath, x 2. 

(d) Three of the warts, with their numerous small spines, x 16. 

(e) Radular teeth. Rhachidian, first, sixth and outermost lateral teeth. Two of the last 
are turned on their sides to show the bulbous basal plates, x 80. 



secretes abundant supplies of a milky white substance from the opaline gland, 
with a strong odour of musk. Extreme irritation, as noted by Mazzarelli (1889 : 582) 
and Engel (1957 : 241) may tinge this secretion with purple, but it is not clear 
whether the colour comes from one of the secreting glands or from the skin. The 
species is a clumsy swimmer. 



REVISION OF THE WORLD SPECIES OF APLYSIA 359 

Head broad, neck short and thick, but both head and neck are narrower than the 
foot and the head projects forwards in front of the foot. Cephalic tentacles short 
and stout, rolled at the edges, linked by rounded lobes to the sides of the mouth. 
Eyes plain. Rhinophores widely spaced, conical, slit almost to the base. 

Foot broad, 35 mm. wide in a specimen 120 mm. long, rounded and winged 
anteriorly, short and rounded posteriorly, in some specimens forming a distinct 
sucker (Text-fig. 41a) (cf. Juliana, p. 365). Penis (Text-fig. 41c) broad, black, in a 
swollen recurved basal portion of the sheath (Text-fig. 416), which is anchored 
by two stout retractor muscles and has a lining covered with wart-like swellings 
armed with spines (Text-fig. 41^). Parapodia widely spaced at their origins, close 
behind the rhinophores, rounded, smooth edged, not large, joined high up posteriorly, 
shutting in the mantle cavity. 

Mantle with oval or round rayed aperture to the shell cavity, and short broad 
siphon. Mantle glands secrete a white substance. 

Mantle cavity closed behind, ctenidium of moderate size, opaline gland large, 
reniform, pigmented, composed of simple vesicles each opening separately to the 
mantle cavity. Genital aperture pigmented, plain or with its edge curled in a spiral, 
hardly covered by the mantle, but well within the protection of the parapodia. 
Genital groove deep, with overhanging dorsal lip. 

Shell 38 x 25 x 3 mm. in a 120 mm. specimen, narrowly ovate, beautifully 
shaped, firm, clear, with concentric and radiating ridges. Hardly any turn-over at 
the apex, but a recurved edge dorsally. Anal sinus wholly posterior, deep and short, 
with a large outer angle. A strong calcareous layer is present in the fresh condition. 

Jaws small, crescentic or nearly rectangular, 9x3 mm. Rods of moderate 
length, straight and rounded at the tips. 

Radula (Text-fig. 410) large, up to 80 rows, formula 40.1.40. Rhachidian tooth 
pale, feeble, with short, high basal plate, straight posteriorly and only slightly 
incised anteriorly. Cusp very short and rounded, with a few indistinct irregular 
denticles. First lateral with an almost triangular plate, broad head, very short obtuse 
cusp and a few small irregular denticles, and one large lateral denticle. Remaining 
laterals with narrowed but still triangular plate, broad head, longer cusp with 
irregular denticles and larger ones both mesially and laterally. Two very small 
vestigial teeth at the end of the row. A simple type of radula. 

Caecum large, its tip curled inwards like a crozier and only the stalk exposed on 
the surface of the digestive gland. 

Cerebral ganglia small, fused. Visceral ganglia distinct, with large granular 
nerve cells. 

Specific Characters 

Large Aplysias, with low broad shape, smooth skin, dark brown or greenish brown 
colour with white or grey blotches and fine veining in black. Secrete a milk-white 
fluid with a strong odour of musk. Foot broad, with a rounded sucker posteriorly. 
Penis stout, black, the bulbous portion of its sheath armed internally with spiny 
warts. Parapodia joined high posteriorly, forming a wall round the mantle cavity. 
Opaline gland large, reniform, pigmented, simple, multiporous. Shell with broad 



3 6o REVISION OF THE WORLD SPECIES OF APLYSIA 

anal angle and posterior anal sinus. Radula with many rows and numerous teeth 
in a row, simple, with few denticles. Caecum crozier-shaped. 
East Atlantic from France to West Africa ; Mediterranean. 



References {see also synonyms) 
Blochmann, 1884 : 32. 
Bohadsch, 1761 : 49, as Lernaea. 
Bosc, 1802 : 63, as Laplysia. 
delle Chiaje, 1828 : 28, 41, 72. 
Grigg, 1949 : 795. 
Mazzarelli, 1891 : 3 and 36. 
Perrier and Fischer, 1908 : 1335. 
Pilsbry, 1895 : 69, as Tethys. 
Rang, 1828 : 64. 
Vayssiere, 1885 : 54, 65 ; 1935, No. 20 (not paginated). 



Aplysia (Aplysia) dura sp. no v. 

Text-figs. 42 and 43 

Distribution. Tristan da Cunha, South Atlantic Ocean, Cook Strait, New 
Zealand. 

Material examined. Two specimens from the inshore waters of Tristan da 
Cunha, taken in the summer of 1952 by Mr. H. F. I. Elliott, and presented by him 
to the British Museum (Natural History). The larger specimen, chosen as the holo- 
type, was given the number B.M. (N.H.), 1957. 6. 11. 1. 

A third specimen came from Lyall Bay, Cook Strait, New Zealand, and is now in 
Wellington Museum, New Zealand, after examination by me. 

The type specimen measures 130 mm. long, 54 mm. wide and 54 mm. high ; 
it is thus of medium size, low and elongated (Text-fig. 42) . Its proportions are unusual, 
for the broad head and long neck are followed by a small mantle and shell area and 
very small low parapodia. The head, including the cephalic tentacles, is 45 mm. 
wide, the rhinophores are 17 mm. apart and are placed mid-way between the cephalic 
tentacles and the commencement of the parapodia, which are 30 mm. apart at their 
origin. The mantle is 45 mm. long, 30 mm. wide, the foot 106 x 40 mm. Black 
pigment, partly bleached, is present on the tentacles, the outer sides of the parapodia, 
the genital aperture and groove, and a little is left on the ctenidium. The inner 
sides of the parapodia are blotched with black, the foot is intense black. There is 
no trace of colour other than the black pigment. Skin very tough and hard, but 
smooth, without a wrinkle. 

Head broad, projecting well in front of the foot. Cephalic tentacles small, rolled 
on the edges. Eyes widely spaced on clear skin areas. Rhinophores wide apart, 
short, stout when contracted. Neck of the same width as the head. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



36i 



Foot broad, tough, rounded in front, with a round sucker posteriorly, tail short, 
projecting behind the parapodial fusion. Penis (Text-fig. 436) very large, stout, 
smooth, black. Bulbous portion of the penis sheath with two strong retractor 





Fig. 42. Aplysia dura. Lateral and dorsal views. 



X 4 . 



muscles and lined with rounded, much branched wart-like swellings bearing spines. 
The warts (Text-fig. 43, b, c) are arranged in rows parallel with the sperm groove, 
the rows occurring in sets of from two to three rows, separated by smooth areas. 
On the side furthest from the sperm groove the warts are scattered and they are 
massed around the base of the penis itself. The warts are elaborately branched, 
with a number of spines on each branch. Parapodia small, thick, short and low, 
but possibly much contracted. They have little flexibility, and are joined posteriorly 



ZOOL. 5, IO. 



19 



362 



REVISION OF THE WORLD SPECIES OF APLYSIA 



forming a mantle cavity pocket, but they do not cover the anal siphon, mantle or 
ctenidium. 

Mantle very small, sharply pointed anteriorly, without overhang here. Mantle 
foramen small, flat, with black radiating lines. Anal siphon short, broad, flat, 
non-tubular. 




R. 



+ 1 



+ 27 



"^Qi^^w 



Fig. 43. Aplysia dura. 

(a) Penis sheath and retractor muscles. X 1 J. 

(b) Penis sheath opened to show the seminal groove, penis and nodosities of the distal 
part of the sheath, x 2. 

(c) A group of the branched nodosities with their numerous stout spines, x 16. 

(d) Shell in dorsal view, x § . 

(e) Radular teeth. Rhachidian and first, 27th and outermost laterals, x 80. 

Mantle cavity small, closed behind. Opaline gland small, simple, hidden beneath 
the muscles of the mantle cavity floor, multiporous. Genital aperture exposed 
(i.e. not covered by the mantle), large, inflated, spiral or smooth. 

Shell (Text-fig. 43d) 40 X 26 mm., narrow and flat. Horny layer thin, with 
concentric lines ; calcareous layer dense, thrown into folds on the edges. No spire, 
anal sinus short and shallow, with the usual broad angle laterally. 

Jaws long and narrow, 15 x 4*5 mm. Rods short, curved. 

Radula (Text-fig. 43^) large and powerful, 20 x 13 mm., at least 90 rows and 



REVISION OF THE WORLD SPECIES OF APLYSIA 363 

formula up to 50.1.50. The teeth are simple, but the denticulations are more 
pronounced than in Juliana. The last four vestigial teeth are very small. 

Salivary glands wide and flat. Caecum large, inflated at the apex, crozier-shaped. 

Cerebral ganglia fused to form a flat band, even the pleuro-pedal mass fused. 

Specific Characters 

The species belongs to the subgenus Aplysia, as shown by the low, broad shape, 
broad foot, fused parapodia, broad siphon, simple opaline gland, spiny warts on 
the penis sheath, short broad pigmented penis, shape of the shell, simple denticula- 
tions of the radular teeth, broad salivary glands, crozier-shaped caecum. But the 
tough but smooth leathery skin, elongated shape, small mantle, shell and visceral 
regions, non-tubular anal siphon, weak opaline gland, more elaborate warts on 
the lining of the penis sheath, and more strongly denticulate, though still simple 
radula, are specific. 

South Atlantic ; South Pacific. 

The species resembles A. cedrosensis from Lower California in its shape, non- 
tubular siphon, small mantle and shell, but differs in the smoothness of its skin 
and in its relatively small visceral region. 

Aplysia (Aplysia) Juliana Quoy & Gaimard, 1832 : 309 

Text-figs. 44, 45 and 46 a, d, e. 

Synonymy : badistes Pilsbry, 195 1 : 1. 

x bipes Pease, i860 : 23, as Syphonota. 

capensis O'Donoghue, 1929 : 14, as Tethys. 

Juliana var. quoy ana Engels & Eales, 1957 : 86, fig. 16a. 

Pparva Pruvot-Fol, 1953 : 38 (see p. 273). 

petiti? Risbec, 1929 : 56. 

rangiana d'Orbigny, 1837 : 2I °- 

sandvichensis Sowerby, 1869 : pi. 4. 

sibogae Bergh, 1905 : 4 ; Baba, 1937 : 211 ; 1949 : 24. 

woodii Bergh, 1908 : 12. 

Rang's sorex (1828 : 57) is a species dubia, as the specimen in the Paris National 
Museum does not agree with the description, and is probably not the type (see 
Engel & Eales, 1957) ; but specimens identified by various authors as sorex are 
usually juveniles of Juliana. 

Distribution. World-wide in warm seas. Western Atlantic from Florida to 
Brazil, including the West Indies ; Eastern Atlantic : Morocco, Ghana and the 
Canary Islands ; Indian Ocean : the Seychelles, Mauritius, Madagascar, Kenya 
to the Cape, India and Pakistan ; in the Western Pacific : Japan, China, Formosa, 
the East Indies, Polynesia, Australia, Tasmania and New Zealand ; Eastern Pacific : 

1 The only species of Pease which can be identified with certainty. 



364 



REVISION OF THE WORLD SPECIES OF APLYSIA 



California, Galapagos and Peru. There is one record from the Mediterranean, the 
Naturhistorisches Museum of Vienna having three specimens from the harbour of 
Algiers. 

Material examined. Numerous specimens from all parts of the world. 

A. Juliana is the most variable species in the genus, and appears to be undergoing 
marked changes in size, form and colour, without anatomical distinctions large 

a) 




(a) 

(b) 

to 
(d) 

to 



X I 



Fig. 44. A ply sia Juliana. 

Lateral view of a specimen from Tasmania, var. bipes. 

Posterior end of the foot to show the sucker, x ij. 

Dorsal view of a specimen from New South Wales, x § . 

Head of this specimen in ventral view, x 1. 

Lateral view of a young specimen from Grand Canary to show partial 



curling up in a 



ball " and the characteristic pinching in of the parapodia anterior to the mantle. 



enough to warrant separation into different species at present. Because of its wide 
range, round the whole world in tropical and sub-tropical seas, one would expect 
to find variations, but it is remarkable that more than one variation occurs in the 
same area. The species was founded by Quoy & Gaimard (1832) from specimens 
taken in the Indian Ocean and figured by them. 

Medium to large, bulky Aplysias, the largest reaching 300 mm. when alive and 
more than half this when preserved (Text-fig. 44, a-e). The general shape is low and 



REVISION OF THE WORLD SPECIES OF APLYSIA 365 

broad with the visceral hump set far back. Colour very variable — yellowish green, 
reddish yellow, dark green, olive, brown, ash grey, or sooty black. There may be 
markings on the sides of the body and the inner sides of the parapodia, mantle 
and mantle cavity floor, tentacles and foot. The edges of the foot may be violet in 
colour. There are usually fine black reticulations or the markings may be darker 
than the ground colour and quite irregular, sometimes in the form of ragged brown 
or black spots, with large and small white flecks and stripes. The skin is smooth, 
but may roughen on contraction or form patterns similar to those seen on leather. 
The contracted animal may assume the sitting hare position and draw in the head 
considerably, or it may curl up in a ball, a condition more commonly seen in juveniles. 
Macnae, in a personal communication, notes the peculiar mode of progression of the 
living animal, which may glide along the surface, or loop along like a caterpillar, 
fastening itself to the substratum by adhesion of the anterior and posterior portions 
of the foot, where the chief pedal glands are situated. In fact the posterior pedal 
glands are especially well developed, as in other members of the subgenus, and the 
foot contracts in this region to form a rounded or heart-shaped sucker. Upon this 
character Mme. Pruvot-Fol founded the subgenus Tullia, but did not realize that 
it is also true of A . depilans, the type of the genus. The sucker, however, is evident 
only when the glands are in a state of secretion, and in a given batch of specimens, 
some will show it well, others feebly and some not at all. The animal has a strong 
smell of musk and when handled exudes copious quantities of an opalescent white 
fluid from the opaline gland on the floor of the mantle cavity. Macnae kindly 
checked this for me on South African specimens. There is no purple secretion from 
the mantle glands. 

G. P. Whitley of the Australian Museum, Sydney, in notes dated June 1924, 
described the egg strings as light to dark yellow, and measured a string of 865 J 
inches, or more than 72 feet. 

Head broad, low and clumsy, with a long but strongly contractile neck. Cephalic 
tentacles flattened, rolled at the edges, quite separate dorsally but joined ventrally 
to each side of the mouth slit. Eyes larger than usual, but may be hidden in con- 
tracted animals. Rhinophores stout, rather small, round when contracted, wide 
apart. 

Foot broad, with rounded edges anteriorly ; in a well expanded specimen there 
may be lateral wings. Posteriorly there is the sucker above mentioned. The penis 
sheath (Text-fig. 450) is large, heavily pigmented, and divided into two portions, 
of which the lower or distal part is bent back on the proximal part, and is attached 
laterally by two strong retractor muscles. The proximal part is muscular, ridged 
internally, with a marked continuation of the spermatic groove. The distal or 
bulbous part is thinner and is lined by numerous simple and compound ingrowths 
or warts, each bearing on its summit small curved spines, which appear to be of a 
chitinous nature (Text-fig. 45&). Young specimens have the warts but no spines. 
The arrangement of the warts varies ; in some specimens they are scattered, in 
the variety bipes they are in rows and groups of rows. They may also spread along 
the penis itself, around its base or along the side not occupied by the spermatic 
groove. A specimen from Formosa had a single row of warts reaching almost to 



3 66 



REVISION OF THE WORLD SPECIES OF APLYSIA 



the tip of the penis. The penis (Text-fig. 45c) is short, dark, very broad and conical ; 
the spermatic groove runs straight along it to near the spoon-shaped tip. Parapodia 
large, evenly rounded, thick and fleshy, freely mobile, natatory, joined posteriorly 
high up, forming a deep posterior mantle cavity pocket. Normally the dorsal area 
is widely exposed, but the parapodia can close over the mantle region. 

Mantle large, thin, about half the length of the contracted animal. Overhang 
wide, covering the large ctenidium. Shell foramen oval, smooth walled, flat, con- 
tractile, sometimes ringed with pigment, rarely rayed. Anal siphon short, wide and 





Fig. 45. Aplysia Juliana. Penis and penis sheath. 

(a) Penis sheath with its retractor muscles, from a New Zealand specimen. X ij. 

(b) Portion of the wall of the distal portion of the sheath of the same specimen, x 16. 

(c) Penis of an Australian specimen, with spiny warts clustered round its base, x 2 J. 



thin, tubular and sometimes fluted along the edges. Mantle glands present, but the 
fluid secreted is white, not purple. 

Mantle cavity extensive, with a large pigmented ctenidium and kidney shaped, 
grey, simple, multiporous opaline gland. Genital aperture smooth and crescentic, 
usually large, strongly pigmented and becoming inflated in the sexually mature 
animal. It is exposed, i.e. not covered by the mantle. The genital groove is strongly 
marked. 

Shell (Text-fig. 46, a, d) variable according to age and habitat, large for the size 
of the animal, but either broad or narrow. In a specimen from Kurachee (= Karachi) 
the length was 1-2 times the breadth ; in a New Zealand specimen the length was 
1*5 times the breadth, giving broad and narrow shells respectively. In general, 
Pacific specimens tend to be narrower than those from the Indian Ocean. One 
character, however, is more constant. The angle made by the anal sinus and the 
right border of the shell, usually referred to as the anal angle, is almost a right 
angle. The shell is not very concave, has a shallow anal sinus, an incurved apex 
without spire and both horny and calcareous layers. Very young specimens may 
exhibit a spiral apex (Marcus, 1955 : fig. 29). 

The jaws are large curved plates of straight or slightly curved, elongate rods. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



367 



Buccal mass large, radula characteristic (Text-fig. 460). The number of rows 
of teeth is high and may reach more than 70, and the formula 40. 1 .40. The feature 




Fig. 46. A ply sia Juliana (a), (d), (e). Aplysia nigra (b), (c) (/). {a-d) shells in ventral 
view, x § ; (e-f ) radular teeth, x 80. 

(a) Shell of A . Juliana, Australian specimen var. bipes (narrow type) . 

(b) Shell of A. nigra (narrow type), New Zealand specimen. 

(c) Shell of A. nigra (broad type), Australian specimen. 

(d) Shell of A. Juliana, specimen from Kurachee, Pakistan (broad type). 

(e) A. Juliana, Australian specimen, var. bipes. Rhachidian, first, sixth and outermost 
lateral teeth of the radula. 

(/) A. nigra var. delli (New Zealand), rhachidian and first and sixth lateral teeth of the 
radula. 



of the dentition is the simplicity of the serrations on the teeth. Rhachidian tooth 
pale and feeble, with a moderately broad basal plate, narrower and deeply excavated 
anteriorly. The cusp is shorter than the plate and has a few irregular small denticles, 



368 REVISION OF THE WORLD SPECIES OF APLYSIA 

which disappear in older teeth. Laterals dark brown in colour, with short, outwardly- 
turned basal plates and short broad heads, the cusp short and plain, except in very 
young teeth, where it bears fine denticles and a single larger denticle on each side. 
The three outermost teeth are degenerate. 

Salivary glands broad and flat. Caecum unusually long and curved inwards like 
a crozier, or spirally coiled near the tip (Text-fig. 8d). 

Cerebral ganglia contiguous within a connective tissue sheath, visceral ganglia 
more or less distinct. 

The following varieties can be distinguished : 

(a) The typical Juliana is large, bulky and of a grey self colour with or without 
ragged spots. It inhabits muddy estuaries in the inshore period and occurs in all 
areas. 

(b) The var. bipes is narrower and smaller, and occurs on the islands in the Pacific, 
on the coasts of East Africa and the East Indies. The spiny warts on the lining 
of the distal part of the penis sheath are arranged in more regular rows than in the 
typical form. 

Using the shell characters alone, two varieties can be distinguished : 

(c) var. quoyana, with narrow shell (Text-fig. 46a). 

(d) var. sibogae, with broad shell (Text-fig. 46^). 

The last two varieties, however, overlap with those given above. Thus bipes 
is narrow shelled, and the typical Juliana may be either broad or narrow shelled, 
according to locality. 

Specimens from Australia carried the Copepod Strongylopleura pruvoti Monod & 
Dollfuss, 1932 on the ctenidium. This is the only species believed to be parasitic 
that is known to occur upon or within any member of the genus. It has also been 
found on the Australian species A. sydneyensis (see p. 350). Specimens were identified 
by Dr. J. P. Harding and have been deposited in the collections of the British 
Museum (Natural History). 

Specific Characters 

Colour and size variable, sooty black, or lighter, with ragged dark spots. Low, 
bulky body, broad foot with posterior sucker on the tail, used for adhesion during 
looping movements ; thick natatory parapodia joined high up posteriorly ; flat 
oval mantle foramen ; white mantle gland secretion ; white musky secretion of 
the simple, multiporous opaline gland ; large radula with feeble denticulations ; 
broad flat salivary glands and crozier-shaped caecum. The bulbous portion of the 
penis sheath lined with fleshy warts, each bearing several hard spines ; the penis 
pigmented, stout, with straight sperm groove. Shell variable, either broad or narrow, 
but with a near right angle between the anal sinus and the right border. 

Circumglobal, approximately from 42 °N. lat. to 46°S. lat. 

References (see also synonyms) 

Adams, H. and Adams, A., 1858, 2 : 33 as julienna. 

Baba, Hamatani & Hisai, 1956 : 216, as sibogae (breeding habits). 

Eales, 1957 : x 79- 



REVISION OF THE WORLD SPECIES OF APLYSIA 369 

Edmonson, 1933 : 152, as Tethys bipes. 

Engel & Eales, 1957 : 8 3- 

Macnae, 1955 : 237. 

Marcus, 1955 : 15. 

Martens, 1880 : 307. 

Ostergaard, 1950 : 99, as bipes, spawning. 

Pilsbry, 1895 : 91, as bipes ; 108, as Juliana ; 86, as rangiana, all as Tethys. 

Pruvot-Fol, 1933 : 400 ; 1934 : 4 1 \ J 953 : 33> as sorex. 

Aplysia (Aplysia) nigra d'Orbigny, 1837 : 20 9 

Text-fig, 46 b, c, f. 

Synonymy : brunnea Hutton, 1875 : 279. 
hamiltoni Kirk, 1882 : 282. 
hyalina Sowerby, 1869 : pi. 5. 

Distribution. Australia (New South Wales), New Zealand (North and South 
Islands), Brazil, Peru. 

Material examined. Specimens from Australia and New Zealand. Dr. Marcus 
sent valuable information relating to living specimens collected on the Brazilian 
coasts. D'Orbigny gave an excellent figure of the animal, copied by Pilsbry. 

This species comes very close to A. Juliana, but the differences in size, colour, 
shape and certain anatomical characters indicate that, unless information to the 
contrary becomes available, it is advisable to retain d'Orbigny's specific name. 
The mollusc is large and bulky, reaching a length of 250 mm. The body is elevated 
and broad, the visceral hump being typically higher and more centrally placed than 
in Juliana. The skin is tough and wrinkled when contracted, heavily and uniformly 
pigmented in black, brownish black or rich brown, usually without pattern, occasion- 
ally with clear areas appearing as white spots. The head, tentacles, sides of the body, 
foot, part of the mantle roof and mantle cavity floor, inner edges of the parapodia 
and the ctenidium are of an intense black or brown colour, but the pigment may 
fade with preservation and become grey or sooty. 

Head broad, neck short compared with Juliana. Cephalic tentacles broad, folded, 
continued to the ventral side of the mouth slit. Rhinophores thick, strongly con- 
tractile, slits short. Eyes small. Genital groove an intense black. Penis broad 
and spatulate, the enlarged bulbous portion of the sheath with both simple and 
elaborately compound (branched) warts bearing numerous spines. The warts are 
partly scattered and partly in regular rows. 

Foot broad, front edges rounded, tail broad, rounded, forming a sucker. Parapodia 
thick, upstanding, commencing close behind but lateral to the rhinophores, capable 
of closing over the dorsal slit, united high up posteriorly, forming a deep mantle 
cavity pocket. 

Mantle large, with considerable overhang and a small, oval or round aperture, 
contractile, sometimes with radiating lines. Anal siphon short and broad. Mantle 



37o REVISION OF THE WORLD SPECIES OF APLYSIA 

glands colourless ; d'Orbigny stated that it exuded a slightly violet and milky 
white secretion, Marcus (1956 : 43) described a black exudate, the local name of the 
animal being " tinteiro " or ink-well. 

Mantle cavity extensive and deep, containing the large, thick, black ctenidium. 
The opaline gland varies from small to large and is involved in the muscles and fibres 
of the body wall ; as usual in the subgenus it is multiporous and consists of simple 
gland cells. D'Orbigny referred to the strong odour of musk from the opaline gland. 
The genital aperture is inflated in the mature animal and becomes thick and fimbriated. 

Shell large, broad and flat, measuring 44 x 37 mm. in a specimen 100 mm. long. 
Anal sinus wide but shallow, apex oblique, dorsal edge recurved posteriorly, anal 
angle almost a right angle but more rounded than in Juliana. 

Jaws composed of very tightly packed, curved, parallel-sided rods, some trun- 
cated and others pointed at the tips. 

Radula large, with numerous rows and many teeth in a row. One specimen 
measuring 131 mm. when preserved had a radular formula of 100 x 60.1.60, 
a figure higher than that of any A. Juliana examined, and with the teeth simpler, 
but with longer cusps to the laterals. The rhachidian tooth is feeble, in some speci- 
mens with hardly a central cusp, and without denticles such as the young tooth 
of Juliana has. Laterals also without trace of denticles except for the single one on 
each side of the rather long, spiky cusp, which at about the middle of a row attains 
i| times the length of the plate. The outermost four teeth are degenerate, three of 
them especially so. 

Cerebral ganglia fused, visceral ganglia large, separate. 

The following varieties can be distinguished : 

[a) var. brunnea resembles nigra, but is deep brown or brownish black all over. 
It has been recorded from New Zealand and Brazil. 

(b) var. delli nov. Specimens from North Island, New Zealand, sent by Dr. R. K. 
Dell of Wellington, had an extension of the parapodial lobes downwards to the foot 
anteriorly, a lower fusion of the parapodia posteriorly, a papillate mantle aperture, 
a tall and leaf -like anal siphon and spiral genital aperture. 

Specific Characters 

Large size, heavy pigmentation in brown or black, usually without pattern but 
occasionally with clear areas on the sides of the body. Mantle aperture small, 
contractile, sometimes rayed. Mantle glands exuding white and black secretion ; 
opaline gland a milky, odoriferous, mucous substance. Shell broad, rounded. Radula 
very large, with a high formula reaching 100 x 60.1.60, the teeth simpler than in 
A . Juliana but with longer and more spike-like cusps to the laterals at the middle 
of the rows. 

Australia, New Zealand, Brazil, Peru. 

References {see also synonyms) 
Allan, 1932a : 423 ; 1950 : 213, as hyalina. 
Marcus, 1956 : 43, as Juliana. 

Pilsbry, 1895 : 85, as Tethys nigra ; 97, as T. brunnea ; 99, as T. hamiltoni. 
Suter, 1913 : 545 and 1915 : pi. 23, as brunnea. 



REVISION OF THE WORLD SPECIES OF APLYSIA 371 

Aplysia (Aplysia) vaccaria Winkler, 1955 : 5, pis. 2, 3 

Text-figs. 47-49 

The holotype is in the Allan Hancock Foundation, University of Southern Cali- 
fornia, No. 983. 

Distribution. Point Fermin, San Pedro, Palos Verdes and San Juan Capistrano, 
Southern California. 

Material examined. A specimen from Doheny Beach State Park, San Juan 
Capistrano, California, sent by Dr. Winkler, and the gut, penis sheath and penis of 
another specimen from the same area also presented by him. 

This large and bulky species was described from living specimens by Winkler 
(1955 : 5), who kept them alive in an aquarium, where they grazed cow-like on the 
seaweed Egregia (Text-fig. 4yd). Alive they attained a length of 255 mm., a width 
of 135 mm. and a height of no mm., with a foot 100 mm. wide. Preserved, the 
specimen now in the British Museum (Natural History) measures 142 mm. long, 
88 mm. wide and 65 mm. high, the foot being no mm. long and 62 mm. wide. The 
rhinophores of the living specimen were 40 mm. posterior to the cephalic tentacles 
and 20 mm. high. The parapodia were 60 mm. apart in front, and formed a wall 
50 mm. high around the mantle cavity posteriorly. The body is stout, firm and 
smooth, of a deep purplish black colour, sometimes without markings, or with 
fine grey to white spots on the sides, head and parapodia. The foot sole is deep 
blue black. There is no odour. The species spawns in February and March, under 
rocks in shallow water, and is usually nocturnal in habit. 

Head large, neck broad. Cephalic tentacles broad, folded, black, fimbriated. 
Rhinophores slender. 

Foot broad, tough, with a short pointed tail and no trace of a sucker. Penis 
sheath (Text -fig. 48a) very large, bent back on itself, fastened by two stout retractor 
muscles attached to the bulbous portion, the black interior of which is lined for 
about half its width by branched swellings capped by curved spines, usually from 
three to five spines on each branch. A few small warts lie on the penis base (Text- 
fig. 48, b, c). Sperm groove white. Penis large, 18 mm. broad at the base, tapering 
to a fine point and curved. Genital groove running straight along the convex side 
of the curve nearly to the tip. Parapodia well developed, widely spaced in front, 
united behind high up, shutting in the mantle cavity. Some mottling on the inner 
sides but no definite pattern. 

Mantle firm, with an aperture 10 mm. wide. Anal siphon tubular, crested, broad. 
Mantle glands secrete a sparse white secretion. 

Mantle cavity closed behind, ctenidium striped black and white, opaline gland 
simple, with many apertures. 



372 



REVISION OF THE WORLD SPECIES OF APLYSIA 



Shell (Text-fig. 47a) large, broad, 70 x 60 X 25 mm. in the holotype, rounded, rather 
thick, with both horny and calcareous layers, no spire but thickened edges fanning 
out from the apex. Anal sinus very shallow and short, the anal angle rounded but 
nearly rectangular. In the B.M. (N.H.) specimen the shell (Text-fig. 47, b, c) measures 
60 mm. long and 43 mm. wide ; it is shallower, but is of the typical Aplysia shape. 





e) 




Fig. 47. Aplysia vaccaria. 

(a) Shell in ventral view. After Winkler, x J. 

(b) Shell of B.M. (N.H.) specimen in dorsal view, x f . 

(c) The same shell in ventral view, x f . In addition to the inturned apex, there is a 
thickening of the chitinous portion along the rim of the shallow anal sinus. 

(d) The mollusc resting on the floor of an aquarium, raising the head as when browsing. 
After Winkler, x f . 

(e) Caecum. 



X 1 



Jaws broad and rectangular, the rods tightly packed, short, straight, truncate 
or rounded at the tips. 

Radula large, with at least 100 rows of teeth and a formula reaching So. 1.80, 
but may be less than this (Text-fig. 49). Rhachidian tooth pale, sunken, feeble, 
with narrow head and simple cusp hardly as long as the basal plate. First lateral 
with tapering, curved basal plate, rounded head and short cusp, bearing a short 
stout denticle on each side. The basal plates broaden and may become squared 



REVISION OF THE WORLD SPECIES OF APLYSIA 



373 



off further along the row, but are never very large. The heads are at first broad 
and rounded, then narrow. The cusps elongate considerably and become narrower, 
reaching their greatest length at about the middle of the row. They are longer than 
in any other species of the subgenus. Their denticles are weak throughout. About 
three teeth at the end of the row are pale, feeble and cuspless. 

Salivary glands very broad and flat. Caecum long, the tip curved inwards in a 
spiral (Text-fig. 470). 




Fig. 48. Aplysia vaccaria. 

(a) Penis sheath and retractor muscles, x f . 

(b) Penis sheath opened to show, conventionally, the warted lining of its basal portion 
and the large, stout, pointed penis. A few warts extend along the convex side of the 
penis. The sperm groove is indicated by a double line ; it runs up the convex side of the 
penis to near its tip. x f . 

(c) A portion of the sheath wall to show the branched warts and spines, x 10. 

The nerve ganglia exhibit complete fusion of the pairs, even the buccal ganglia 
are fused. 



Specific Characters 

Large, bulky Aplysias, with smooth skin, black coloration, broad dark foot with 
pointed tail on which no sucker has been observed. Penis large, black, in a bulbous 
sheath lined by branched spiny warts. Parapodia ample, joined high up behind. 
Mantle aperture large, flat, mantle glands secrete a white substance. Anal siphon 
tubular. Opaline gland simple, multiporous. Shell broad, thick, with shallow anal 
sinus. Radula large, with numerous rows and teeth in a row, all the teeth simple, 
but with cusps of considerable length. Caecum long, with spirally coiled apex. 



374 REVISION OF THE WORLD SPECIES OF APLYSIA 

Pairs of ganglia fused. The characters are those of subgen. Aplysia, except for the 
absence of a sucker on the foot. 
Southern California. 

R +1 +6 + 24 +35 




Fig. 49. Aplysia vaccaria. Radular teeth. Rhachidian and first, sixth, 
24th and 35th laterals, x 100. 

The species has affinities with A. cedrosensis, but the latter has a thick, warted 
skin, brown mottled coloration, flat, non-tubular anal siphon, simple warts on the 
penis sheath lining, and shorter cusps to the lateral teeth of the radula. 



Subgenus Phycophila A. Adams, 1861 

Aplysia (Phycophila) euchlora (Adams in M. E. Gray, 1850) 

Text-fig. 50 

Synonymy : Placobranchus euchlorus Adams in M. E. Gray, 1850, 2 : tab. 179 fig. 1, 
from a drawing by A. Adams ; 4 : 35 (Explanation of plates). 

" Aclesia euclorus (Pacobranchus e.), Adams '^ M. E. Gray op. 
cit. 4 : 98. 

Aplysia (Phycophila) euchlora A. Adams, 1861 : 41. 

Aplysia immunda Bergh, 1902 : 163 (juvenile). 

Distribution. Western Pacific : Japan (Adams's specimen) and Siam (Bergh's 
specimen and one in the B.M. (N.H.)). 

Material examined. A single specimen from Siam (Text-fig. 50, a-e), label- 
led A. mouhoti Gilchrist, 1895, M. Mouhot's collection, numbered B.M. (N.H.), 
i860. 1. 18. 3. This specimen is not A. mouhoti, which is a synonym of Syphonota 
geographica. 

1 The mis-spellings are in this section only of Mrs. Gray's work. The type-species of Placobranchus 
Hasselt is a nudibranch. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



375 



A. Adams gave a figure and a very imperfect account of his species. It was green, 
smooth, compressed, with narrow sole, elongated anterior tentacles, narrow rhino- 
phores truncate at their apices, and long slender tail. Shell membranous, oblong, 
dilated in front, the apex not involute. It was found in the Strait of Tsugaru, Japan, 
crawling on floating weed. Bergh's specimen was a juvenile 17 mm. long, 9 mm. 



a) 




b) 





T 



d) 



+6 



+ 12 



*Mh SA ft *w 

Fig. 50. Aplysia euchlora. 

(a) Dorsal view, x 1 . 

(b) Head in ventral view, x 1 . 

(c) Shell in dorsal and ventral views, x §. 

(d) Jaw rods. 

(e) Radular teeth, Rhachidian, first, sixth, 12th and outermost laterals, x 80. 

wide and 8 mm. high, elongated, on floating algae, from Siam. He described a net- 
work on the sides, with scanty black spots, and a median black band dorsally. Sole 
of the foot narrow, only 5 mm. wide. Inner sides of parapodia with strong black 
perpendicular bands joined at intervals. Mantle glands carmine. Radula 34 x 

This meagre information on the species is hardly sufficient for identification, were 
it not for certain peculiar features. These are the extreme narrowness of the body 
and especially of the foot, the unusually long slender tail, and the posterior narrowing 
of the parapodia in Adams's figure. The British Museum specimen agrees so well 



376 REVISION OF THE WORLD SPECIES OF APLYSIA 

with these features, and in its locality, Siam, that it must either be identified as 
euchlora or made into a new species. The following account is based on the assump- 
tion that it belongs to Adams's species. 

The specimen (Text-fig. 50a) measures 65 mm. long, 20 mm. wide and 24 mm. 
high. It is soft and flabby, but not much contracted. There are remnants of black 
pigment on the sides and foot, but no definite markings. There are a few black 
marks on the inner sides of the parapodia, a little pigment on the ctenidium and the 
cephalic tentacles, but the rhinophores are colourless. 

Head and neck narrow, the latter short. Cephalic tentacles (Text-fig. 50&) broad, 
hardly rolled on the edges but turned over along the anterior border, where they 
extend across the head anterior to the mouth, with a small central depression ; 
laterally they are pointed and slope backwards. The rhinophores are short, close 
together, diverging, slender. Eyes rather large, anteriorly placed relative to the 
rhinophores. 

Foot narrow, only 7 mm. wide, very soft and wrinkled, with round anterior 
border and long, pointed tail. Penis sheath small, not very muscular, its distal end 
recurved, with two small retractor muscles. Penis filiform, very long and narrow. 
Parapodia moderate, very thin on the edges, open widely exposing the mantle 
region, free and flexible, coming close together posteriorly over the foot, but hardly 
joined. 

Mantle small, 18 X 15 mm., thin, delicate, without visible foramen or colour 
in the mantle gland. Anal siphon small, thin, projecting backwards in the specimen. 

Mantle cavity exposed, ctenidium small, opaline gland reniform, simple, with 
many apertures. Genital aperture in front of the mantle, small, crescentic, genital 
groove feeble. 

Shell (Text-fig. 50c) 26 x 18 mm., very delicate, oval, rather flat, with very small 
apex, anal sinus long, shallow. Remains of calcareous layer present. 

Jaws small, very dark brown. Rods of exceptional length, closely and neatly 
packed, giving a tesselated appearance, some spatulate, others near the edge curved 
at the tips like birds' heads (Text-fig. 50^) . 

Radula (Text-fig. 50*2) of dense brown colour but broken up. Buccal mass small. 
At least 40 rows and a formula 25 . 1 . 25. Rhachidian tooth with broad base, slightly 
curved behind and excavated in front, cusp at first longer than the basal plate, 
but shortening with wear, denticulations of the usual type in the genus. Laterals 
with short curved basal plates, tapering but becoming bulbous near the end of the 
row, with long, denticulate cusps. The last three or four teeth vestigial but unusually 
dark in colour. 

Cerebral ganglia fused into a flat band, visceral ganglia joined. 

Specific Characters 

Small size, slender shape, very narrow foot and long pointed tail. Cephalic tentacles 
broad and of unusual shape, rhinophores slender, close together, eyes more anteriorly 
placed than usual. Penis filiform. Parapodia free, meeting on the tail. Mantle 
small, shell sac closed. Opaline gland simple, multiporous, genital aperture not 
covered by the mantle. Shell oval, flat, delicate, with long, shallow anal sinus. 



REVISION OF THE WORLD SPECIES OF APLYSIA 377 

Radula without special characteristics, but small and dark in colour. Cerebral 
ganglia fused. Habitat on floating weed. 
North Pacific Asiatic area. 



3. GEOGRAPHICAL DISTRIBUTION 

Distribution of the species of Aplysia is limited by three factors — the life history 
of the mollusc, its feeding habits and its relation to temperature. Aplysia feeds on 
seaweed and is therefore confined to the littoral and sub-littoral zones of the land 
masses and islands of the world. There is some evidence that it is an annual. The 
eggs in vast numbers are laid in small circular capsules fastened together in long 
strings and deposited between tide marks (see p. 295). The free-swimming larvae 
proceed to open water, settling after metamorphosis in the red weed zone on the 
sea bottom, where they feed and grow and eventually migrate shorewards to the 
brown weed zone, changing colour from red to brown. They remain in shallow 
water on the brown weed (Laminaria in this country) until mature, when they 
migrate again to the Fucus zone, between tide marks. Here there may be many 
thousands at a time, but only for a short period. They couple in chains of from three 
to ten or even more, the top member of the chain acting as male only, the lowermost 
member as female only, and all the intermediate ones as male for the lower and female 
for the upper individual in the chain. After spawning, which ensues shortly after 
separation from coupling, many die, but it is believed that some return to deeper 
water. Inshore migration and spawning may occur twice in the year or only once 
(see Eales, 1921 : fig. 5). Thus distribution is affected by the presence of suitable 
weed and spawning grounds. 

Approximately two-thirds of the known species inhabit tropical and sub-tropical 
waters, three at least (dactylomela, Juliana and parvula) being circumglobal. Only 
one (punctata) has been recorded from within the Arctic Circle, and none is known 
so far from the Antarctic. 

A study of the distribution of species in all the oceans suggests a division of the 
world into ten regions. This division does not always follow the most obvious 
boundaries of the oceans (Text-fig. 51). 

1. North Atlantic, west, comprising the east coast of North America and Central 
America, Bermuda and the West Indies. A. dactylomela, Juliana and parvula are 
common, particularly from Florida southwards, also brasiliana and cervina, while 
north of 40 north latitude niorio and willcoxi reach Rhode Island and New England 
respectively, although mono also extends further south. 

2. North Atlantic, north-east, consisting of the coasts of Europe and the Mediter- 
ranean. Three species are typical of this area. A. depilans and fasciata range from 
the Atlantic coasts of France, Spain and Portugal to become the dominant species 
in the Mediterranean. Although both have occasionally appeared on the British 
coasts, they do not extend northwards, but have spread southwards along the 
African coasts and the islands (see 3 below) . In the north the only species is punctata, 
which reaches Greenland, Norway and the Baltic and is the only truly indigenous 

ZOOL. 5, 10. 20 



37» 



REVISION OF THE WORLD SPECIES OF APLYSIA 




REVISION OF THE WORLD SPECIES OF APLYSIA 379 

British species. Southwards its range is the Mediterranean and the islands off 
Africa. 

Recently three specimens from the Harbour of Algiers, now in the Naturhistorisches 
Museum in Vienna, proved to be young Juliana, the first record of that species in 
this area. 

3. North Atlantic, south-east, including the west coast of Africa from Morocco 
to the Gulf of Guinea, with the Azores, Madeira, the Canary Islands and Cape Verde 
Islands. A. punctata has been recorded from the Canaries, depilans from the coasts 
of Morocco, Rio de Oro and Madeira, and fasciata all down the coast to Senegal 
and Ghana. A. dactylomela, Juliana and parvula occur both on the islands and the 
mainland, brasiliana is occasionally taken in Ghana, but winneba is one of the com- 
monest species after dactylomela. 

4. South Atlantic, west, comprising the east coasts of South America from Guiana 
to Cape Horn. The southern part of this area is little known, most of the records 
and specimens coming from Brazil. Here brasiliana, cervina, dactylomela, Juliana, 
nigra and parvula are found, although cervina is typically a more northerly species. 

5. South Atlantic, east, that is, the west coast of Africa from the Gulf of Guinea 
to the Cape, and the islands of St. Helena and Tristan da Cunha. This region also 
is imperfectly known. A. brasiliana has been found on St. Helena, fasciata as far 
south as Angola and the tough species dura on Tristan da Cunha. The east-west 
drift of ocean currents from South-west Africa towards Brazil may account for the 
distribution of brasiliana. 

6. Indian Ocean, west. This extends from the Red Sea and the Persian Gulf 
in the north, down the east coast of Africa to the Cape, and includes the islands of 
the Seychelles, Mauritius, Reunion, and Madagascar, and the coasts of Pakistan, 
India and Ceylon. The area is rich in species and good collections have been made. 
The three circumglobal species dactylomela, Juliana and parvula are common, the 
first penetrating the Red Sea from the south, while fasciata has reached it from the 
north ; oculifera occurs all round the coasts of the Indian Ocean, even to the west 
coast of Australia, but maculata is confined to the west, from Mauritius to South 
Africa. A. cornigera is a common species in India and Ceylon, gracilis inhabits the 
Red Sea. 

7. Pacific Ocean, north-west, comprising the eastern and south-eastern coasts of 
Asia, Japan, Formosa, the Philippines and East Indian Islands. Again an area 
rich in species and inhabited by dactylomela, Juliana and parvula. A. cornigera 
occurs in the Philippines and in Indo-China, oculifera in the East Indies, Formosa 
and Japan. Arthur Adams recorded a number of species from Japan, including 
euchlora, the floating species. To Bab a we owe the species kurodai and sagamiana 
from Japan. A South Pacific species which extends northwards to the Asiatic 
continent is pulmonica. 

8. North Pacific, north-east, comprising the west coast of North and Central 
America. Here live some of the giant members of the genus, such as californica 
of California and the Gulf, cedrosensis, and rehderi of the Gulf, alongside the ubiqui- 
tous dactylomela, Juliana and parvula. The large black vaccaria, also of the subgenus 
Aplysia, occurs on the coast of south California, robertsi in Central America. 



380 REVISION OF THE WORLD SPECIES OF APLYSIA 

9. South Pacific, west, including the whole of Australia for convenience, Tasmania, 
New Zealand and Polynesia. This region has been well worked and is rich in species. 
The three circumglobal species are present, Juliana exhibiting varieties of consider- 
able range, from the small spotted bipes var. to larger and broader forms. In Australia 
in addition to parvula, dactylomela, Juliana and nigra the characteristic species 
are cronullae, denisoni, gigantea, reticulata, sowerbyi and sydneyensis, with oculifera 
in western Australia and extraordinaria in the east. New Zealand has, in addition 
to the three world-wide species, keraudreni, a large and handsome species described 
by Rang from Polynesia, while in North Island dura (see Zone 5) has been found. 
In Polynesia the fauna of the Hawaiian Islands is best known. Again the three 
circumglobal species are found, and in addition extraordinaria and Gould's pulmonica. 

10. South Pacific, east, comprising the west coasts of South America and the 
Galapagos Islands. This region is little known, most of our information being 
obtained from specimens collected in Peru during the early part of the nineteenth 
century, by d'Orbigny, Eydoux and Souleyet. A. Juliana occurs on Galapagos 
and d'Orbigny's nigra came from Peru. His inca from the same coast is a valid 
species, but Rang's lessoni is probably keraudreni. Mazzarelli & Zuccardi's chierchiana 
is doubtful. 

It will be noted that while some species have an unlimited range within the tropics 
and sub-tropics, others are local, even when mixed with the ubiquitous species. 
No explanation can be offered of this, and it is probable that more extensive collecting 
would extend the range of the presumed local species. On the whole, distribution 
of most species can be accounted for by proximity of islands to land masses, e.g. 
Madeira and the coast of West Africa. Currents explain the distribution of brasiliana 
at St. Helena, Ghana and in the western tropical Atlantic ; and the presence of 
dura at Tristan da Cunha and New Zealand is paralleled in the distribution of 
fishes (see Regan, 1913). The scarcity of species on the west side of South America 
may be due to the steepness of the shores and depth of the water near the coast. 

4. CONCLUSIONS 

The subdivision of the genus into the five subgenera Pruvotaplysia, Neaplysia, 
Varria, Aplysia and Phycophila (see p. 271) indicates that the genus is undergoing 
change and radiation, from the primitive parvula, through the majority of the more 
generalized members, to the extremes in Aplysia and Phycophila. The subgenus 
Neaplysia is a small offshoot, including, it is true, very large members of a single 
species, but their only deviation from the main group (subgen. Varria) is a peculiarity 
of that already vestigial structure, the shell. The floating members of the subgenus 
Phycophila are little known. 

The primitive characters of Pruvotaplysia, with its two species, parvula and 
punctata, are the high fusion of the parapodia posteriorly, closing in the mantle 
cavity behind, the large, flat, non-contractile mantle aperture, the deeply concave 
shell, small simple radula and the separate rounded nerve ganglia. 

Varria contains about 70% of the known species, and exhibits the greatest genera- 
lization in structure, together with a wide range in distribution. In the posterior 



REVISION OF THE WORLD SPECIES OF APLYSIA 



38i 



Distribution of the Species of Aplysia According to the Zones in Section 3, p. 377 

Zones 





r 














— ^ 


Species 


1 


2 


3 


4 5 


5 7 


8 


9 


10 


brasiliana 


X 




X 


X X 










californica 










. 


X 






cedrosensis 












X 






cervina 


X 






X 










cornigera 










>< X 








cronullae 














X 




dactylomela . 


X 




X 


x . . : 


< X 


X 


X 




denisoni 










X 




X 




depilans 




X 


X 




. . 








dura 








X 






X 




euchlora 


. . 








X 








extraor dinar ia 


. . 








. 




y 




fasciata 




X 


X 




X 








gigantea 










. 




X 




gracilis 










X 








inca 
















X 


Juliana 


X 


X 


X 


X 


X X 


X 


X 


X 


keraudreni 














X 


X 


kurodai 










X 








maculata 










X 








morio . 


X 
















nigra 








X 






X 


X 


oculifera 










X X 








parvula 


X 




X 


X 


X X 


X 


X 


X 


pulmonica 










X 




X 




punctata 


. 


X 


X 












rehderi . 












X 






reticulata 


, . . 












X 




robertsi 












X 






sagamiana 










X 








sowerbyi 


. 












X 




sydneyensis . 














X 




vaccaria 












X 


. . 




willcoxi 


X 












. . 




winneba 






X 













spout of the parapodia and flexible mobile anal siphon, contractile, occasionally 
closed, mantle aperture, and fusion of the cerebral ganglia and the visceral pair, the 
subgenus links Aplysia with Syphonota. 

Rang used the mantle aperture for classificatory purposes, but although in Pruvot- 
aplysia and Aplysia it can be relied upon, in Varria it varies with age, state of con- 
traction and even with individuals. Thus a young animal may possess an aperture 
which normally closes in the adult, an adult may show an open aperture or a closed 
contracted one, with or without a papilla at the point of closure. Occasionally the 
closed aperture may have a fringed edge. These variations make the mantle aperture 
unreliable for identification of the species. 



382 REVISION OF THE WORLD SPECIES OF APLYSIA 

In Varria, brasiliana resembles an enlarged, more slender and more active cervina, 
with narrower foot and more elongated head ; pulmonica is like a plainer and flatter 
dactylomela, without rings on the skin ; winneba resembles a smaller and broader 
fasciata, yet there appears to be no doubt that they are distinct, and their areas 
overlap. A. sagamiana is of interest because the discrete nature of its nerve ganglia 
recalls that of Pruvotaplysia, although in all other characters its relationships are 
with Varria. 

The members of the subgenus Aplysia exhibit the greatest deviation from the 
characters shown by the majority of the species, and also the most marked variation 
within one species. The broad foot, usually with temporary formation of a posterior 
sucker, the absence of purple secretion but abundance of a white, milky, odoriferous 
substance from the simple, multiporous opaline gland, the numerous but extremely 
simple teeth of the radula, the spiral caecum, and the large, stout penis lying in 
a sheath armed with spiny warts, are peculiar to the subgenus, which contains six 
representatives. 

In Phycophila the body is adapted for floating on algae ; it is narrow and stream- 
lined, with a long pointed tail resembling that of Stylocheilus . Unfortunately there is 
no information on its breeding habits ; it would be interesting to know whether it 
comes inshore to lay its eggs, or whether the floating habit is permanent and the egg 
ribbons are coiled round the weed. 

Aplysia is by far the largest genus in the family Aplysiidae, containing at least 
35 species. Pruvotaplysia is more primitive than any other section of the family ; 
Varria has characters which resemble those of Syphonota and to a lesser extent the 
Dolabellinae ; the subgenus Aplysia is the most specialized and links the Aplysiinae 
with the Dolabriferinae and the Notarchinae because of its armed penis sheath. 



5. ACKNOWLEDGMENTS 

It is with pleasure that I thank the British Museum (Natural History) Trustees 
for giving me the privilege of working through their collections of Aplysia and for 
a grant in aid, and especially for the practical help, always freely given, of Mr. 
I. C. J. Galbraith, M.A., Curator of Molluscs, Mr. H. O. Ricketts and the late Mr. 
G. L. Wilkins. 

Museum authorities and their staffs have been most generous in lending their 
material for identification and investigation, and my thanks are due to : 

Dr. R. K. Dell of the Dominion Museum, Wellington, New Zealand. 

Dr. W. K. Emerson of the American Museum of Natural History, New York 
City. 

Professor Dr. H. Engel of the Zoologisch Museum, Amsterdam. 

Dr. H. Lemche of the Universitets Zoologiske Museum, Kobenhavn. 

Dr. D. McMichael of the Australian Museum, Sydney. 

Miss Hope Macpherson of the National Museum of Victoria, Melbourne. 

Dr. Nils H. Odhner of the Naturhistoriska Riksmuseet, Stockholm. 

Dr. Oliver E. Paget of the Naturhistorisches Museum, Wien. 

Dr. R. L. C. Pilgrim of Canterbury University College, Christchurch, New Zealand. 



REVISION OF THE WORLD SPECIES OF APLYSIA 383 

Prof. Dr. Gilbert Ranson of the Museum National d'Histoire Naturelle, Paris. 

Dr. Harald A. Rehder of the Smithsonian Institution, Washington, D.C., U.S.A. 

Dr. R. Tucker Abbott of the Academy of Natural Sciences, Philadelphia, U.S.A. 

To the following I am grateful for recently collected material, and /or notes on 
the living animals, sketches and colour photographs : — 

Mr. R. Bassindale of the Zoology Department, University of Bristol, for West 
African specimens. 

Dr. Elizabeth Batham of Portobello Marine Biological Station, University of 
Otago, New Zealand. 

Dr. T. Habe of Amakusa Marine Biological Laboratory, Kyushu, Japan. 

Dr. R. B. Hill, Marine Biological Station, St. George's, Bermuda. 

Dr. Alison Kay of the Department of Zoology, University of Hawaii, Honolulu. 

Dr. H. Lemche of Copenhagen, University Museum. 

Dr. William Macnae, formerly of Rhodes University, Grahamstown, now at the 
University of the Witwatersrand, Johannesburg. 

Dr. D. McMichael of the Australian Museum, Sydney. 

Professor R. D. Purchon of the University of Malaya in Singapore. 

To Dr. Ernst and Frau Eveline Marcus, of the University of Sao Paulo, Brazil, 
I express my thanks for descriptions of South American species and for answers to 
my numerous questions. Dr. L. R. Winkler, of the School of Tropical and Preventive 
Medicine, Loma Linda, California, sent me specimens and collaborated with me 
regarding the diagnosis of the species A. vaccaria described by him in 1955. 



6. SPECIES REPRESENTED IN MUSEUM COLLECTIONS 

A list of the species represented in the various museums is appended : 

British Museum (Natural History). The complete list, with localities, is given for reference. 

Aplysia brasiliana Rang, 1828 

Specimens 

1856. 1 1. 21. 31 No loc. Mr. Brandt's coll. Purchd. .... 1 

1868.4.8. 1 St. Helena. Pres. J. C. Mellis, Esq 1 

1869.6.5.74 Rio Janeiro Bay. 5 fms., 1867. Pres. Lords of the 1 

Admiralty (coll. Dr. Cunningham) 

1 958. 1. 9. 1-2 Gold Coast, W. Africa. Dr. F. R. Irvine ... 2 

Aplysia calif ornica Cooper, 1863 

1958 .1.9. 3-4. Bird Rock, La Jolla, Calif., 1954. Below tide marks. Coll. 2 

Miss A. M. Clark 
1958. 1. 9. 170-3. Doheny Beach State Park, Calif. 7. v. 1957. From Dr. 4 

L. R. Winkler. Pres. Dr. N. B. Eales 

Aplysia cervina Dall & Simpson, 1902 

1924. 11 .24.4. Pernambuco, Brazil. Pres. Eastern & Assoc. Tel. Co . 1 

1958. 1. 9. 5. Pian Prince. Zool. Soc. coll. ..... 1 



3^4 



REVISION OF THE WORLD SPECIES OF APLYSIA 



1958. I. 9.6-10. 



Aplysia cornigera Sowerby, 1869 
Ennur (E.), 1937. Krusadai. Winckworth coll. 



Specimens 
5 



1958. 1. 9. 11. 

1958. 1. 9. 12. 



Aplysia cronullae Eales 

Cronulla, Sydney, N.S.W. From Dr. H. Lemche. Pres. 

Dr. N. B. Eales . . . . . Paratype 

New South Wales. From Sydney Nat. Mus. 12083. Pres. 

Dr. N. B. Eales 



1958.1.9.13-14- 

1846. 8. 31. 3. 
1868.7.4.63. 

1876.2.5.9. 
1876.2.5. 10. 
1890. 12.28.93-4. 
1901.7.20.17. 
1913.10.10.1. 

1927. 11. 2. 2. 
1932. 11. 23. 1-2. 
1954. 12. 10. 1-3. 
1958. .1.9. 15 
1958. 1. 9. 1 6-20. 



1958. 1. 9 
1958. 1. 9 

1958. 1. 9 

1958. 1. 9 

1958. 1. 9 

1958. 1. 9 
1958. 1. 9 

1958. 1. 9 
1958. 1. 9 
1958. 1. 9 

1958-1.9 
1958. 1. 9 
1958. 1. 9 
1958. 1. 9 
1958. 1. 9 
1958. 1. 9 
1958. 1. 9 



21-3. 

24. 

25- 
26. 

27. 

28. 

29-30. 

31-2. 

33- 

34-6- 

37- 

186-7. 

188. 

195-199- 
200. 
201. 
245- 



Aplysia dactylomela Rang, 1828 

St. Vincent, W.I. Rev. L. Guilding (re-registered from 
1839.12.27.30) . ... 

Jamaica. Coll. Mr. Gosse. Purchd. Mr. Cuming 

St. Vincent. Pres. Lords of the Admiralty (coll. Dr 
Cunningham) ...... 

Mauritius. Purchd. Mr. Cutter 

Shore reefs, Tonga. J. J. Lister . 

Kingston Harbour, Jamaica. J. E. Duerden, Esq 

Found on beach at Mosquito Bay, St. Thomas, W.I. Pres 

Capt. A. Morrel .... 

Tavinni, Fiji. Dr. H. S. Evans 
Little Abaco, Bahamas. L. R. Crawshay 
Schoenmaker's Kop, Easter, 1952. Pres. Dr. Wm. Macnae 
Anse Corail, Mahe. 22.xii.38. Winckworth coll 

20.X.37. 22.xii.38. Winckworth 

coll ... 

Mahe, Seychelles. Winckworth coll. 

S. E. Cay, Port Royal, Jamaica. 26.viii. 1939. Camb. U 

Jamaica Exp. 1939 (J. S. Colman) 
Salt Isl., Portland, Jamaica. i3.viii. 1939. Camb. U 

Jamaica Exp. 1939 (J. S. Colman) 
Sta. Colon 4 ? Panama Canal. Sci. Exp. Res. Ass. Pacific 

Cruise 1923-24. C. Crossland .... 
St. Kitts, B.W.I. Coll. and pres. A. W. Vaughan, 1951 
Takoradi, Gold Coast. Pres. Mrs. Carlyle Bell 
Accra, Gold Coast. Irvine & Foote 
No loc. No coll. or donor 
„ ,, H. Welton, Esq. 
No coll. or donor 



Miss Alison Kay. Pres. Dr. N. B. Eales 



Kanai, Hawaii 

Honolulu. 

Pram Pram, Gold Coast 



1958. 10. 10.2. 



4.11.1950 
,, ,, ,, ,, 21. xi. 1949. 

i7- i - I 949- 
Winneba shore, Gold Coast. 22 . xi . 1949 

dale ....... 

West Shore, St. George, Bermuda, 28. vi. 1958 

Dr. R. B. Hill. Pres. Dr. N. B. Eales 



Pres. R. Bassindale 



Pres R. Bassin- 



Coll. by 



REVISION OF THE WORLD SPECIES OF APLYSIA 



385 



Specimens 
1958.10. 10.3-4. Parham Sound, Antigua. Pres. Dr. J. W. Gregory . . 2 

1958.12.30.94. Long reef, N.S. Wales (F 19935) Victoria Nat. Mus. 

Melbourne, via Dr. N. B. Eales .... 1 



1881 . 11 .10.11. 
1881.11 .10.3. 
1923. 1.26.6-9. 



1928.7.27.4 

1958. 1. 9. 38 

1958. 1. 9. 39 

1958. 1. 9. 40 

1958. 1. 9. 41 



Aplysia denisoni Smith, 1884 

Port Denison, Queensland. Pres. Lords of the Admiralty 
(coll. Dr. Cunningham) .... Holotype 

Port Molle and Port Denison, Queensland. Pres. Lords of 
the Admiralty (coll. Dr. Cunningham) 

Abrolhos Isls. Pres. W. J. Dakin & C. H. O'Donoghue 
(Dakin's Abrolhos coll.) 

Cocos Keeling I si. Pres. Col. I. St. J. Grant 

Sydney. J. B. Jukes, Esq. 

No loc. ,, ,, 

No coll. or donor 



1845.9.26. 13. 
1848.4.27. 13. 

1850.6. 17. 12. 
1856. 1. 18. 6. 

i860. 1 1. 28. 1. 

1866.7.30. 13. 
1958. 1. 9. 165-7. 

1898. 5. 21. 109-10. 
1958. 1. 9. 164. 
1870. 1. 14. 20. 

1958. 1. 9. 193. 



Aplysia depilans Linn., 1791 

S. of Europe. Pres. P. B. Webb, Esq. . 

R. Tweed. Pres. Dr. G. Johnston . 

Weymouth Bay. Mr. Damon's coll. 

Mediterranean ? Zool. Soc. coll. (sent to Soc. by Mme 

Jeannette Power) ...... 

R. Itchen, Southampton. Pres. J. S. Davies, Esq. . 

Adriatic. Purchd. L. W. Schaufuss 

North Coast of Spain and Portugal. " Noma " Exp., 1870 

Coll. Mr. Kent. Purchd. Mr. Geale (re-registered from 

1872.2.3. 124) 

Zool. Stat. Naples. Norman coll. .... 
No loc. No coll. or donor. (As A. camelus Cuv.) . 
Polperro, coast of Cornwall. Pres. J. Couch, Esq. (Holo 

type of A. melanopus Couch, 1870) 
Marseilles. Peterson, Esq. (Holotype of A. petersonii 

Gray, 1828) . . . . . 



1957. 6. 11. 1. 

1957. 6. 11. 2. 



Aplysia dura Eales 

Inshore waters Tristan da Cunha. S. Atlantic. Summer 

1952. Coll. and pres. H. F. I. Elliot . . Holotype 

Inshore waters Tristan da Cunha. S. Atlantic. Summer 

1952. Coll. and pres. H. F. I. Elliot . . Paratype 



i860. 1. 18. 3. 



Aplysia euchlora A. Adams, 1861 
Siam. M. Mouhot's coll. (as A. mouhoti Gilchrist, 1895) . 



1883. 11. 12. 1-3. 

1958. 1. 9. 42. 

1958. 1. 9. 43. 



Aplysia extraordinaria Allan, 1932 

Port Jackson. From N.S. Wales Coast Internat. Fisheries 

Exhib., 1883 

No loc. No coll. or donor (old coll.) .... 



3 86 



REVISION OF THE WORLD SPECIES OF APLYSIA 



1958. 1. 9. 194. 
1958. 1. 9. 168-9. 



1873 
1898 
1908 
1958 
1958 

1958 
1958 

1958 
1958 

1958 

1958 



31. 1. 

21.125. 
22.1-2. 
9.44. 

9-45- 

9.46. 
9.47-8. 

9.49. 
9.252. 

9.253. 

9.387-8. 



Aplysia fasciata Poiret, 1789 

Mediterranean ? Zool. Soc. coll. (sent to Soc. by Mme 

Jeannette Power) (re-registered from 1856. 1. 18. 6) 
North coast of Spain and Portugal. " Noma " Exp. 1870. 

Coll. Mr. Kent. Purchd. Mr. Geale (re-registered from 

1872.2.3. 124) . 
Benguela, Africa. Purchd. Mr. Monteiro 
Zool. Stat. Naples. Norman coll. .... 
Las Palmas, Grand Canary. Dr. J. Cleasby Taylor 
Christiansborg, Gold Coast, 9.x. 36. Dr. F. R. Irvine 

Castle rocks, Gold Coast. 12.iii.1950. Dr 

F. R. Irvine 

Accra, Gold Coast. Irvine & Foote 

Arrecife, Lanzarote Isl., Canary Isls. Coll. and pres. Capt 

A. K. Totton 

Marseilles. Baron G. Cuvier .... 

Chorkor shore, Accra, Gold Coast. 12. hi. 1950. Pres. R 

Bassindale ........ 

Trawled 4 fms., 2 miles off shore, Accra. 2. hi. 49. 

Pres. R. Bassindale ...... 

St. I. Madeira. 25.lv. 24. "St. George" Exp., 1924 

C. Crossland ....... 



Specimens 



1860.4. 14.32. 
1923.1.26.10-11. 



Aplysia gigantea Sowerby, 1869 

Fremantle, W. Australia. Purchd. Dr. Bowerbank 

Pres. W. J. Dakin & C. H. 
O'Donoghue (Dakin's Abrolhos coll.) . 



Aplysia gracilis Eales 

1907 . 1 1 . 15 . 29. Nr. Gemil, Lake Menzaleh, Egypt. 

of Education, Govt, of Egypt 



Pres. His Excell. Min. 
Holotype 



1908.9.22.3-4. 
1958. 1. 9. 50-1. 

1958. 1.9. 52-3. 

1958. 1. 9. 54-9. 

1958. 1. 9. 60-7. 
1958. 1.9.68. 
1958. 1. 9- 69. 

1958. 1. 9- 177. 

1958. 1. 9- 178-9. 
1958. 1. 9. 180. 
1958. 1.9. 190. 



Aplysia Juliana Q. & G., 1832 

Las Palmas, Grand Canary. Dr. J. C. Taylor 

Port Alfred, South Africa. 8. hi. 1955. Pres. Dr. Wm. 

Macnae ......... 

Hermanus, Cape of Good Hope. Jan. 1949. Pres. Dr. 

Wm. Macnae ........ 

Arrecife, Lanzarote Isl., Canary Isls. Mar.-May 1955. 

Coll. and pres. Capt. A. K. Totton .... 
St. I. Madeira. 28. iv. 1924. " St. George " Exped., 1924 
Galapagos. C. Crossland coll. 1923-24. 
Port Phillip, Victoria, Australia. J. B. Wilson coll. (has 

dried up at some time) ...... 

Fishers Isl., New York. Jan. 1950. Coll. H. L. Ferguson. 

Pres. Dr. N. B. Eales * 

Tomioka, Amakusa, Japan. Pres. Dr. N. B. Eales . 



Keelung, Taiwan (Formosa) 
N. B. Eales 



Mr. I. Callan. Pres. Dr. 



REVISION OF THE WORLD SPECIES OF APLYSIA 387 

Specimens 
1958. 1 .9.246. Gold Coast, Africa. Pres. R. Bassindale ... 1 

1958. 1. 9. 389-92. St. I. Madeira. 25.iv.24. "St. George" Exped., 1924. 

C. Crossland ........ 4 

1958.12.30.95. Long Reef, N.S. Wales (F19934) Victoria Nat. Mus., 

Melbourne, via Dr. N. B. Eales ..... 1 

1958.12.30.96. Portland Bay, N.S. Wales (F18980) Victoria Nat. Mus., 

Melbourne, via Dr. N. B. Eales ..... 1 

Aplysia Juliana var. bipes Pease, i860 

1958 .1.9. 70-9. Newcastle, N. H. [New South Wales]. Rev. C. P. N. Wilton 14 

1958. 1.9.80. Peru. Rev. W. Hennah ...... 1 

1958. 1. 9. 81-5. Fort Jesus, Mombasa. 27-30. x. 1952. Pres. L. F. 

Brown ......... 5 

+ egg-mass 
1958. 1. 9. 86-91. Diamond Head Reef Flat, Oahu, Hawaii. 23. xi. 1956. 

Pres. Dr. N. B. Eales 6 

Aplysia Juliana var. quoyana Engles & Eales, 1957 

1881 . 10.6.55. Port Jackson. Pres. Admiralty. Dr. Coppinger, H.M.S. 

" Alert " . . . . . . . Holotype 1 

1883. 11. 12. 4-5. Port Jackson. From Internat. Fisheries Exhib., 1883 . 2 

1887. 12.22. 1-6. Hakodadi, Japan. Pres. Rev. W. Andrews ... 8 

1921.12.20.291-2. Misaki, Japan. A. V. Insole ..... 2 

1958. 1. 9. 95. Port Jackson. J. B. Jukes (1845) .... 1 

Aplysia Juliana var. sibogae Bergh, 1905 

1883.4.25. 19. Kurrachee. Exch. with Kurrachee Mus. 1 

1958. 1. 9. 181. Tomioka, Amakusa, Japan. Pres. Dr. N. B. Eales . . 1 

Aplysia keraudreni Rang, 1828 

1958. 1. 9. 96-7. Channel N. of Quarantine Isl., Otago Harbour. + 2 fms. 

18" dredge. From Portobello Mar. Biol. Sta., Otago 
Harbour, New Zealand. Pres. Dr. N. B. Eales . . 2 

Aplysia kurodai Baba, 1937 

1921. 12.20.293-6. Misaki, Japan. A. V. Insole ..... 4 

1958. 1. 9. 184-5. Tomioka, Amakusa, Japan. Pres. Dr. N. B. Eales . . 2 

Aplysia maculata Rang, 1828 

1954. 12. 10. 14-17. Hermanus, Cape Province. Jan. 1949. Pres. Dr. Wm. 

Macnae ......... 4 

1958. 1. 9. 102-6. Hermanus, Cape Province. Jan. 1949. Pres. Dr. Wm. 

Macnae ......... 5 

Aplysia morio Verrill, 1901 

1958. 1. 9. 107. Florida. U.S. Nat. Mus. Pres. Dr. N. B. Eales . . 1 

1958. 10. 10. 1. Ferry Reach, St. George's West, Bermuda. June 1958. 

Coll. by Dr. R. B. Hill. Pres. Dr. N. B. Eales . . 1 



3 88 



REVISION OF THE WORLD SPECIES OF APLYSIA 



1886. 6. 3.61. 

1958.1.9.93- 
1958. 1. 9. 94. 
1 Q58. 1 2. 30. 97-9. 



Aplysia nigra d'Orbigny, 1837 

Port Jackson. Purchd. Dr. von Lendenfeld . 

No loc. Haslar coll. J. O. Goodsir .... 

New South Wales ....... 

Drummoyne, N.S. Wales (F19931). Victoria Nat. Mus., 
Melbourne, via Dr. N. B. Eales . . . . . 



Specimens 
1 



1886. 11. 18. 3. 



Aplysia nigra var. brunnea Hutton, 1875 

Dunedin, New Zealand. Purchd. Comm. for New Zealand, 
India and Colon. Exhib., 1886 ..... 



1958. 1. 9. 92. 



Aplysia nigra var. delli Eales 



North Isl., New Zealand. 
Dr. N. B. Eales 



From Wellington Mus. Pres. 
Holotype 



1869.7-8.35- 
1883.4.25.20. 
1954. 12. 10. 13. 

1958. 1. 9. 108. 

1958. 1. 9. 109-13. 

1958. 1. 9. 114. 
1958.1.9.191-2. 



1958. 1.9. 115-16. 

1877.4. 16. 14. 
1954. 12. 10. 4-12. 

1954. 12. 10. 18-27. 

1958. 1 .9. 117-18. 
1958. 1 .9.119-20. 
1958. 1. 9. 121-30. 
1958. 1. 9. 1 34-40. 



1958. 1. 9. 131-33. 

1958.1.9-182-3. 
1919. 10.7.55. 

1958. 12.30. 100-1. 



Aplysia oculifera Ad. & Rve., 1850 

Red Sea, Gulf of Suez. R. Mc Andrew . 
Kurrachee. Exch. with Kurrachee Mus. 
Flat rocks beyond Hotel Pollock, Port Elizabeth. Pres 

Dr. Wm. Macnae ..... 
Abu Zabad, on reef at low tide, Aqaba. 1 1 . ii . 1949. Coll 

N. B. Marshall 

Bokness, Alexandria Div., S. Africa. Mar. 1954. P fes 

Dr. Wm. Macnae ..... 
Trincomali. 16. hi. 38. Winck worth coll. 
Reef N. Ngatpaet Passage, E. Babethuap, Palau Isls 

Ostheimer ? Sta. 425. Pres. Acad. Nat. Sci. Philad 

(Tucker Abbott) 

Aplysia parvula (Guilding MS.) Morch, 1863 

St. Vincent, W.I. Purchd. Rev. L. Guilding's sale (re 

registered from 1839.12.27.30) .... 
S. Africa. Sent by Dr. Daniel Hahn ... 
Flat rocks beyond Hotel Pollock, Port Elizabeth. Pres 

Dr. Wm. Macnae ..... 
Rocks at Kariega mouth, C. of Good Hope. Pres. Dr. Wm 

Macnae ....... 

W. Indies. No donor or coll. 

No loc. ,, ,,,,,, 

New South Wales. Godwin-Austen coll. 

Miller's Point, Cape Peninsula, S. Africa. Dec. 1949 

Pres. Dr. Wm. Macnae .... 

Berbera, Brit. Somaliland. Feb. 1949. Pres. A. Fraser 
Brunner ......... 

Tomioka, Amakusa, Japan. Pres. Dr. N. B. Eales . 

S. Cheval Paar, Ceylon. Coll. Prof. Herdman, 1902. Pres. 
T. J. Evans (Holotype of A. intermedia Farran, 1905) 

Portland, Victoria (F18981) Victoria Nat. Mus., Melbourne, 
via Dr. N. B. Eales ....... 



2 

1 

21 

29 
2 
2 

14 



+ egg-mass 



REVISION OF THE WORLD SPECIES OF APLYSIA 



339 



Aplysia pulmonica Gould, 1852 

1848.4.3. 13. Norfolk I si. Sir J. Richardson 

1877. 1 .15. 16. Samoa Isl. Pres. Rev. S. J. Whitmee . 

1958. 1 .9.141-2. Anini, Kauai, Hawaii. 31.xii.1956. Pres. Dr. N. B 

1958. 1. 9. 143. Port Molle. J. B. Jukes .... 

1958. 1. 9. 144. Port Jackson. J. B. Jukes .... 



Eales 



Specimens 
1 

2 
2 

1 
1 



1958. 1. 9. 145. 



Aplysia pulmonica var. tryoniana Pilsbry, 1895 
No loc. J. B. Jukes (1845) .... 



1851.7.4.4. 
1858.5.28.5. 

1862.10.27.33. 
1865. 2. 11. 16. 
1866. 12. 31. 5. 

1868.3.6.62. 
1872.2.3. 164. 

1888. 12. 3. 8-10. 
1898.5.21.111-13. 
1927.2.24.2-3. 
1927.2.24.4-5. 
1931.6.23.8-9. 
1946.7.27. 1. 
1947.6. 12. 1. 
1947.6. 12.7. 

1948.7. 10. 1. 

1958. 1. 9. 146-7. 
1958. 1. 9. 148-9. 

1958. 1. 9. 150. 

1958. 1. 9. 151. 
1958. 1.9. 152-3. 
1958. 1. 9. 154-7. 
1958. 1. 9. 158. 
1958. 1. 9. 159-60. 



Aplysia punctata Cuvier, 1803 

? Tynemouth. Pres. Slater . 
Guernsey. Purchd. Mr. Joshua Alder . 

Bohuslan, Sweden. A. W. Malm, No. 66 

Scotland. W. Carruthers, Esq. 

Taken in crab pots 20 fms., Polperro, Cornwall. Purchd 
Laughrin 

? Europe. Purchd. of Damon 

N. coast of Spain and Portugal. " Noma " Exp. 1870 
Coll. Mr. Kent. Purchd. Mr. Geale 

Firth of Lorn. 17-30 fms. J. Murray 

Zool. Stat. Naples. Norman coll. . 

Kames Bay, Cumbrae. Mrs. Stebbing 

Ilfracombe. Mrs. Stebbing . 
? donor . 

Selsey Bill. Pres. E. Heron- Allen 

Plymouth Sound, 1946. Purchd. Mar. Biol. Ass. Plymouth 

Yealm Estuary, Devon. Purchd. Mar. Biol. Ass. Ply- 
mouth. Coll. W. J. Rees, 8.^.1947 

Ceibach Beach, Newquay, Cardiganshire, Wales. Pres 
Wm. D. Evans 

Marseilles. G. Cuvier . 

Sardinia. Prof. Bonelli 

Devon. G. Montague 

Torbay. Mrs. Griffiths 

Probably British. No donor or coll. 

No loc. ,, ,, ,, ,, 



Aberdaron Bay, N. Wales, 15-20 
Coll. D. D. John & F. C. Fraser . 



fms. 27.vii.1936 



1 

5 

+ spawn 

1 
1 

4 

1 

1 
3 
3 

2 
2 
2 
1 
1 



1886.2.22.29. 
1887.5. 12.3. 



Aplysia reticulata Eales 

N.W. Australia. Purchd. Capt. Beckett. 

Dampier Archipelago, N.W. Australia. 

Beckett ...... 



Holotype 
Purchd. Capt. 



1958. 1. 9. 161, 



Aplysia robertsi Pilsbry, 1895 
N. coast, Central America. Pres. Capt. Dow 



39© 



REVISION OF THE WORLD SPECIES OF APLYSIA 



1958. i. 9. 189. 
1958. 12.30. 102-4. 



Aplysia sowerbyi Pilsbry, 1895 

Specimens 
Long Reef, Sydney, N.S.W. E. Pope and party. Pres. 

Dr. N. B. Eales 1 

Western Port, Victoria (F18984) Victoria Nat. Mus. 
Melbourne, via Dr. N. B. Eales ..... 3 



1883. 11. 12. 6. 



Aplysia Sydney ensis Sowerby, 1869 

Port Jackson. N. S. Wales Coast Internat. 
Exhib. 1883 



Fisheries 



Aplysia vaccaria Winkler, 1955 

1958. 1 .9. 174. Doheny Beach State Park, 2 m. SW. San Juan Capistrano, 

Calif. 15. v. 1957. From Dr. L. R. Winkler. Pres. 
Dr. N. B. Eales 

1958. 1 .9. 175. Digestive gland and penis. From Dr. L. R. Winkler. 

Pres. Dr. N. B. Eales 



1934. 9. 11. 4. 
1958. 1. 9. 176. 



Aplysia willcoxi Heilprin, 1886 

Dry Tortugas. G. Tandy & J. S. Colman 
Fishers Isl., New York. Jan. 1950. Coll. H. L. Ferguson. 
Pres. Dr. N. B. Eales 



I957-5- I4-I- 

I957-5-I4 
I957-5-I4 



1957.6. 18 
1957.6. 18 

1957.6. 18 

1957.6. 18 

1882.8. 16 



3- 

1. 

2-3- 

4-7- 

8. 
17-18. 



1958. 1. 9. 162. 

1958. 1. 9. 163. 
1958. 1 .9.202-11. 

1958. 1. 9. 212. 

1958. 1. 9. 213-19. 

1958. 1. 9. 220-7. 



Aplysia winneba Eales, 1957 

Chorkor, Accra, Gold Coast. 14.x. 1949. Pres. R. Bas- 

sindale ....... Lectotype 1 

Chorkor, Accra, Gold Coast. 14.x. 1949. Pres. R. Bas- 

sindale ..... Paratype (figured) 1 

Chorkor, Accra, Gold Coast. 14.x. 1949. Pres. R. Bas- 

sindale ....... Paratype 1 

Nr. Dixcove, Gold Coast. Pres. Miss V. J. Foote Paratype 1 

Pram Pram, Gold Coast. Aug. 1939. Pres. Miss V. J. 

Foote ....... Paratypes 2 

Christiansborg, nr. Accra, Gold Coast. Pres. Miss V. J. 

Foote ....... Paratypes 4 

Accra, Gold Coast. Pres. Dr. F. R. Irvine & Miss V. J. 

Foote ....... Paratype 1 

Gold Coast. Pres. Maj. Burton & Capt. Cameron 

Paratypes 2 

Takoradi, Gold Coast. Pres. Mrs. Carlyle Bell . . 1 

Santiago Isl., C. Verde. 23.lv. 1939. Pres. C. R. Stoner 1 

Pram Pram, Gold Coast. 4.11.50. Pres. R. Bassindale . 

Paratypes 14 

Pram Pram, Gold Coast. 4.11.50. Pres. R. Bassindale 

Paratype 1 

Pram Pram, Gold Coast. 21.xi.49. Pres. R. Bassindale 

Paratypes 7 

Winneba shore, Gold Coast. 15.xi.49. Pres. R. Bassin- 
dale ....... Paratypes 8 



REVISION OF THE WORLD SPECIES OF APLYSIA 391 

Specimens 
1958 .1.9. 228. Axim Hospital Reef, Accra, Gold Coast. 13 . iv . 49. Pres. 

R. Bassindale ...... Paratype 1 

1958. 1.9.229-30. Chorkor, Gold Coast. March 1949. Pres. R. Bassindale 

Paratypes 2 

1958. 1. 9. 23 1-6. Dredge haul No. 2, Gold Coast. 4.1.51. Pres. R. Bas- 

sindale ....... Paratypes 6 

1958. 1 .9.237-40. Christiansborg, Gold Coast. 15.1.49 and 17. hi. 49. Pres. 

R. Bassindale ...... Paratypes 4 

1958. 1 .9.241-4. Dixcove shore, Gold Coast. 7.11.51. Pres. R. Bassindale 

Paratypes 4 

1958. 1. 9. 247-50. Gold Coast. Pres. R. Bassindale . . . Paratypes 4 

1958. 1 .9.251. Chorkor shore, Gold Coast. 12. hi. 50. Pres. R. Bassin- 

dale ....... Paratype 1 



Types of nominal species not now included in the genus Aplysia 



Syphonota geographica Ad. & Rve. 

1882.2.23.574-5. Thursday Isl., Torres Straits, 4-5 fms. H.M.S. "Alert". 

Coll. Dr. Coppinger. (Holotype and Paratype of 
A. piperata Smith, 1884.) 

1882. 2. 23. 571-3. Thursday Isl., Torres Straits, 4-5 fms. H.M.S. "Alert". 

Coll. Dr. Coppinger. (Holotype and 2 Paratypes of 
A. sparsinotata Smith, 1884.) 

i860. 1. 18. 2. Siam. M. Mouhot's coll. (Holotype of A. Mouhoti Gil- 

christ, 1895) 



Australian Museum, Sydney, New South Wales : dactylomela, extraordinaria (holotype), 
Juliana, nigra, parvula, reticulata, sydneyensis, sowerbyi. 

Australia, National Museum of Victoria, Melbourne : dactylomela, extraordinaria, Juliana, 
oculifera, parvula, sydneyensis. 

New Zealand, Christ Church Museum, Otago and Portobello Marine Biological Station : Juliana, 
keraudreni, nigra, parvula. 

New Zealand, Dominion Museum, Wellington : dura, Juliana, nigra, keraudreni, parvula. 

United States National Museum (the Smithsonian Institution), Washington, D.C., U.S.A. : 
brasiliana, calif ornica, cedrosensis (holotype), cervina, dactylomela, Juliana, morio, parvula, 
pulmonica, punctata, rehderi (holotype), willcoxi. 

Philadelphia, Academy of Natural Sciences : brasiliana, californica, cervina, dactylomela, 
depilans, Juliana, oculifera, parvula, punctata, robertsi, willcoxi. 

Stockholm, Naturhistoriska Riksmuseum : brasiliana, californica, cervina, dactylomela, depilans, 
morio, parvula, pulmonica, sagamiana. 

Kjabenhavn Universitets Zoologiske Museum : brasiliana, cervina, cornigera, cronullae (type), 
dactylomela, fasciata, gigantea (shell), Juliana, keraudreni, morio, oculifera, parvula, pul- 
monica, punctata, sydneyensis. 

Paris, Museum National d'Histoire Naturelle : brasiliana, californica, cervina, cornigera, dacty- 
lomela, depilans, extraordinaria, fasciata, inca (type), Juliana, kurodai, maculata, oculifera, 
parvula, pulmonica, punctata. 

Wien, Naturhistorische Museum : brasiliana, californica (shells), dactylomela, fasciata, Juliana, 
kuyodai, morio, oculifera, parvula, sydneyensis. 



392 



REVISION OF THE WORLD SPECIES OF APLYSIA 



7. REFERENCE LETTERS 
abd.ao. — abdominal aorta. mant. — mantle. 

an. — anus. mant. ap. — mantle aperture. 

ant. giz. — anterior portion of gizzard. mant. cav. — mantle cavity. 

ant. giz. t. — anterior gizzard tooth. mant. f. — fold of mantle beginning to en- 

an. s. — anal siphon. velope the shell. 



an. si. — anal sinus in shell. 
apex. — apex of the shell. 

b. pi. — basal plate of tooth. 
bl. — blade of tooth. 

caec. — caecum. 

calc. — calcareous portion of shell. 

ceph. t. — cephalic tentacle. 

ct. — ctenidium. 

cu. — cusp of tooth. 

d.— denticle of tooth. 

e. — eye. 

/.—foot. 

g. buc. — buccal ganglion. 
g. c. — cerebral ganglion. 
g. par.— parietal ganglion. 
g. ped. pedal ganglion. 
g. pi. — pleural ganglion. 
g. v. — visceral ganglion. 
gen. ap. — genital aperture. 
gl. herm. hermaphrodite gland. 

haem . — haemoco ele . 
hd. — head of tooth. 

int. — intestine. 



op. gl. — opaline gland. 

op. gl. ap. — aperture of opaline gland. 

p. giz. — posterior portion of gizzard. 
p. gl. — purple gland. 

pap. — papilla on mantle. 

para . — parapodium . 

para. sin. — parapodial sinus. 

pc. — pericardium. 

ped. a. — pedal artery. 

ped. gl. — pedal gland. 

ped. lac. — pedal lacuna. 

ped. n. 3. — third pedal nerve. 

ped. su. — pedal sucker. 

pn. — penis. 

pn. s. — penis sheath. 

pz. — sensory (brush) cell. 

R. — rhachidian tooth. 

r. bd. — right bile duct. 

r. pi. vise. conn. — right pleuro-visceral con- 
nective. 

rec. — recurved border of shell. 

ren. p. — renal pore. 

ren. s. — renal sac (kidney). 

ret. mant. — retractor muscle of visceral 
hump. 

ret. pn. — retractor penis muscle. 

rh . — rhinophore . 



/. ped. m. — longitudinal pedal muscle. 
I. pi. vise. conn. — left pleuro-visceral 

nective. 
la. herm. d. — large hermaphrodite duct. 
lat. — lateral tooth, 
liv. — digestive gland. 



sem. gr. — seminal groove, 
con- sh. — shell. 

sh. f. — shell-forming region of the mantle. 
stom. — stomach. 

t. — tail. 



8. LITERATURE 
Adams, A. 1861. On some new species of Mollusca from the north of China and Japan. 

Ann. Mag. nat. Hist. ser. 3, 8 : 135-142. 
Adams, H. & Adams, A. 1853-58. The genera of recent Mollusca arranged according to their 

organisation. 2 vols : London. (Vol. 2 includes Aplysia.) 
Adams, A. & Reeve, L. 1850. The Zoology of the Voyage of H. M.S. Samarang 1843-46. In 

three parts. London, 1848-50 : i-x, 1-24 ; 25-44 > 44~88 ; pi. 1-24. (Part 3 contains 

Aplysiids.) 



REVISION OF THE WORLD SPECIES OF APLYSIA 393 

Aelian, C. c. a.d. 150. De natura animalium. Lib. 2 and Lib. 16. 

Allan, J. K. 1932a. Sea hares. Austr. Mus. Mag. Sydney, 4, 12 : 419-425. 

1932&. A new genus and species of sea slug and two new species of sea hares from 

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1950. Australian shells, pp. 1-470. Georgian House, Melbourne. 

Angas, G. F. 1867. A list of species of Marine Mollusca found in Port Jackson Harbour, 
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1877. A further list of additional species of Marine Mollusca to be included in the fauna 

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Arango y Molina, R. 1878. Contribucion a la Fauna Malacologia Cubana. An. R. Acad. 

de Ciencias, Habana, 1-280 -j- 1-35. 
Baba, K. 1935- A general sketch of the Molluscs inhabiting the Coral reef of the Okinawa 

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1936. Opisthobranchia of the Ryukyu (Okinawa) Islands. /. Dep. Agric. Kyusyu 

Univ. 5, 1 : 1-49. 

1937a. Opisthobranchia of Japan, 1. Ibid. 5, 4 : 195-236. 

1937b. Tethys and Aplysia, with a synopsis of Tethys from Japan. Zool. Mag., Tokyo, 

49, 2 : 57-63. 

1938. Opisthobranchia of Kii, Middle Japan. /. Dep. Agric. Kyushu Univ. 6, 2 : 1-19. 

1949. Opisthobranchia of Sagami Bay collected by His Majesty the Emperor of Japan. 

Tokyo : 1-104. English translation : 1 21-194. 

1955- Ibid. Supplement : 1-59. Tokyo. 

Baba, K. & Hamatani, I. 1952. List of the species of the Opisthobranchia from Kii, Middle 

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Wiesbaden, 1897-1902, 7 Bd. 5 Abth. Anaspidea, 313-382. Also 9 Bd. 6 Abth. 1908 : 
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1823a. Art. " Lievre marin ", ibid. 26 : 316-328. 

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96 : 277-288. 

1825. Manuel de Malacologie et de Conchy liologie. Paris : 1-472. Plates 1827. 

Blochmann, F. 1884. Die im Golfe von Neapel vorkommenden Aplysien. Mitt. zool. Stat. 

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Bosc, L. A. G. 1802. Histoire naturelle des Vers. 1 : 1-324. 
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Proc. linn. Soc. N.S.W. 2 : 74-89. 

zool. 5, 10. 21 



394 REVISION OF THE WORLD SPECIES OF APLYSIA 

Burn, R. 1957. On some Opisthobranchia from Victoria. /. malac. Soc. Aust. 1 : 11-29. 
Burne, R. H. 1906. Notes on the anatomy of South African Aplysiidae, with description of 

two new species. Proc. malac. Soc. Lond. 7 : 51-58. 
Cantraine, F. 1840. Malacologie mediterraneene et littorale. Mem. Acad. R. Bruxelles, 

13 : 1-71. 
Carus, J. V. 1889-93. Prodromus Faunae Mediterraneae. 2, 1 : 1-854. 
Chiaje, S. delle. 1822-29. Memorie sulla storia e notomia degli animali senza vertebre del 

regno di Napoli. 4 vols. Naples. (Aplysia Vol. 1, 1822, 25-76.) 

1828. Decrizione e Notomia delle Aplisie. Atti R. Instit. Incoragg. Napoli, 4 : 25-76. 

Clessin, S. 1899. Die Familie de Aplysiidae, in Systematisches Conchylien Cabinet von Martini 

und Chemnitz. Niirnberg. 1, 8 : 1-56. 
Cockerell, T. D. A. 1901. A new Tethys from California. Nautilus, 15, No. 8 : 90-91. 

1915. Tethys (Neaplysia) californica (Cooper). Ibid. 29, No. 7 : 84. 

Colgan, N. 1908. General account of dredging operations. Dublin Marine Biological Com- 
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191 1. Marine Mollusca. Biological survey of Clare Island in the county of Mayo, Ire- 
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■ 1914. The Opisthobranch fauna of the shores and shallow waters of the County Dublin. 

Irish Nat. Aug.-Sept. 1914 : 161-204. 
Cooper, J. G. 1863. On new or rare Mollusca inhabiting the coast of California. Proc. 

Calif. Acad. Sci. 3 : 56-60. 
Cotton, B. C. & Godfrey, F. K. 1933. South Australian Shells. Part 7. 5. Aust. Nat. 

14, 3 : 72-108. 
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son anatomie, et sur quelques-unes de ses especes. Ann. Mus. Hist. nat. Paris, 2 : 287-314. 

1805. Suite de l'extrait des memoires sur les Mollusques. Bull. Sci. Soc. philom., Paris, 

3, No. 93 : 254-256. 

1817a. Le Regne Animal. Gastropodes, 2 : 382-401. 

18176. Memoires pour servir a Vhistoire et V anatomie des Mollusques. 9. Mem. sur le 

genre Aplysia. Paris. 1-28. 

1829. Rapport verbal sur la monographic des Aplysiens de M. Rang, fait a l'Academie 

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Dakin, W. J., Bennett, Isobel & Pope, Elizabeth. 1953. Australian Sea Shores. London, 

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Proc. U.S. nat. Mus. 6 : 318-342. 

1885. List of Marine Mollusca comprising the Quaternary Fossils and Recent forms from 

American localities between Cape Hatteras and Cape Rogue, including the Bermudas. 
Bull. U.S. geol. geogr. Surv. 24 : 1-336. 

Dall, W. H. & Simpson, C. T. 1902. The Mollusca of Porto Rico. Bull. U.S. Fish Comm. 

20, part 1, 1900-01 : 353-516. 
Darwin, C. i860. Journal of Researches into the Natural History and Geology of the Countries 

visited during the voyage of H.M.S. Beagle round the world under the command of Capt. 

FitzRoy, R.N . London. 2nd Edition : 519. 
Dautzenberg, P. 1929. Contribution a la Faune de Madagascar. Mollusca 2, Mollusca 

marina testacea. Faune Colon, franc. 3 : 321-637. 
Dautzenberg, P. & Bouge, J. L. 1933. Les Mollusques testaces marins des etablissements 

francais de l'Oceanie. /. Conchyliol. 11 : 41-108. 
Dautzenberg, P. & Fischer, P. H. 1925. Les mollusques marins du Finistere. Trav. Sta. 

biol. Roscoff, Fasc. 3 : 1-180. 



REVISION OF THE WORLD SPECIES OF APLYSIA 395 

Depontaillier, J. 1880. Note sur YAplysia ? grandis Philippi et YAplysia deperdita Philippi. 

/. Conchyliol., 20, 3 : 251-255. 
Deshayes, G. P. 1839-53. Traite elementaire de Conchy liologie. 2 vols. Texte pp. i-xii, 

1-824 ; 1-384. PL pp. i-xi, 1-80 ; pis. 1-132. Atlas 1864. 

1857. Note sur diff6rents mollusques de la Guadeloupe, envoyes par M. Schramm. J . 

Conchyliol. 6 : 2 ser. t. 2. 137-143. 

1863. Catalogue des Mollusques de l'lle de la Reunion (Bourbon) : in L. Maillard : 

Notes sur Vile de la Reunion. 2, Molluscs : 1-140. 

Dioscorides, P. Born 36 b.c. De medica materia. Lib. 2. Cap. 18. 

Dobson, G. E. 1880. Notes on Aplysia dactylomela. J. Linn. Soc. (Zool.) 15 : 159-160. 

Dufo, H. 1840. Observations sur les Mollusques marins, terrestres et fluviatiles des lies 

Sechelles et des Amirantes. Ann. Sc. nat. {Zool.) (2), 14 : 166-221. 
Eales, N. B. 192 1. Aplysia. Mem. 24, Liverpool Mar. biol. Committee, Liverpool : 1-84. 

1944- Aplysiids from the Indian Ocean, with a review of the family Aplysiidae. Proc. 

malac. Soc. Lond. 26, 1 : 1-22. 

1957a. Aplysiids from West Africa, with description of a new species, Aplysia winneba. 

Ibid. 32, 4 : 179-183. 

19576. Revision of the species of Aplysia of the Museum National d'Histoire Naturelle 

(Malacologie) Paris. Bull. Mus. Hist. nat. Paris. (2), 29, No. 3 : 246-255. 

1957c. Proposed use of the plenary powers to validate the specific name parvula Morch, 

1863, as published in the combination Aplysia parvula (Class Gastropoda). Bull. zool. 
Nom. 13, part 8 : 240-244. See Int. Comm. Zool. Nom. Opin. 560, 1959. 

Edmonson, C. H. 1933. Reef and shore fauna of Hawaii. Honolulu, Hawaii, Spec. Publ. 

Bishop Mus. 22 : 1-295, 163 t. f. 
Ehrenberg, C. G. 1828 (plates) and 1831 (text). Symbolae Physicae Evertebrata. Animalia 

Mollusca. Berlin. (Pages not numbered.) 
Eliot, C. 1899. Notes on Tectibranchs and naked Mollusca from Samoa. Proc. Acad.nat. 

Sci. Philad. (3), 51 : 512-523. 
Engel, H. 1927. Westindische opisthobranchiate Mollusken. 2. Aplysiidae, etc. Bijdr. 

Dierk. 25 : 83-122. 

1929. Aplysia dactylomela Rang, a circumtropic species. Proc. malac. Soc. Lond. 18: 

I47-I5I- 

19340. Are the genera and species of Bohadsch 1761 to be accepted? Ann. Mag. nat. 

Hist. (10), 13 : 529-540. 

19346- Les Aplysies de M. de Blainville. /. Conchyliol. 78 : 83-90. 

1936a. Uber westindische Aplysiidae und Verwandten anderer Gebiete. Capita zool. 

8 (1) : 1-76. 

19366. On the names of the genera Tethys and Aplysia. Temminckia. 116 : 221-266. 

1957- On the influence of preservation on the external appearance of specimens of 

Aplysia depilans L. Beaufortia, 5, No. 66 : 241-243. 

Engel, H. & Eales, N. B. 1957. The species of Aplysia belonging to the subgenus Tullia 

Pruvot-Fol, 1933 : on a generic character in statu nascendi. Ibid. 6, No. 69 : 83-114. 
Farran, G. P. 1905. Report on the Opisthobranchiate Mollusca. In Ceylon Pearl Oyster 

Fisheries Reports, Part 3, Suppl. Rep. 21 : 329-364. Roy. Soc. Lond. 
Ferussac, A. E. 1821-22. Tableaux systematiques des Animaux terrestres ou fluviatiles, vivants 

oufossiles. Mollusques: xlvii, 1-30, 114. Paris and London. 
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nat. St. Servan, 8 : 11-15, and 10 : 25-31. 
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Forbes, E. & Hanley, S. 1853. A History of British Mollusca and their Shells. 3 : 1-616. 
Garstang, W. 1890. A complete list of the opisthobranchiate Mollusca found at Plymouth. 

/. Mar. biol. Ass. U.K., n.s., 1 : 399-457. 
Gilchrist, J. 1895. On a group of the Aplysiidae, with description of a new species. Ann. 

Mag. nat. Hist. (6), 15 : 403-405. 

ZOOL. 5, IO. 2I§ 



396 REVISION OF THE WORLD SPECIES OF APLYSIA 

Gould, A. A. 1852. Mollusca and Shells of the U.S. Exploring Expedition, 1838-42. Phila- 
delphia, 12 : 223-226 (Gastropoda 1-383). 
Gray, J. E. 1828. Spicilegia Zoologica. Pt. 1 : pp. 4-5, pi. 4, figs. 4, 4a. (4 London). 

1847. A list of the genera of recent Mollusca, their synonyma and types. Proc. 

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1857. Guide to the systematic distribution of Mollusca in the British Museum. London. 

Part 1 : 1-230. 

, Mrs. M. E. 1842-57. The figures of molluscous animals . . . for the use of students. 

London. 5 Vols. Parts relating to Aplysia are : pis. 61, 61* and 61** (vol. 1, 1842), 

pis. 138-149, 177, 179 (vol. 2, 1850) and the explanations of these plates in vol. 4 (1850), 

pp. 12, 30-31 ; also list of species in vol. 4 (pp. 97-98). 
Grigg, Ursula M. 1949. The occurrence of British Aplysiae. /. Mar. biol. Ass. U.K. 28 : 

f 795-805. 
Guerin Meneville, F. E. 1844. Iconographie du Regne Animal de G. Cuvier. Mollusques. 

Paris, 3 : 1-64. Planches 1-104. Text dated 1844, plates between 1829 and 1843. 
Guiart, J. 1 90 1. Contribution a l'etude des Gasteropodes Opisthobranches et en particulier 

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its neighbourhood. 1. Molluska. Proc. Sci. Assoc. Trinidad, 2 : 134-157. 

1895. The Molluska of the Gulf of Paria. Proc. Vict. Inst. Trin. 2 : 1 16-152. 

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of Queensland. Rep. 12th Mtg. Aust. Ass. Advanc. Sci. 1909 : 343-371. 

1916. A preliminary index of the Mollusca of Western Australia. /. roy. Soc. W. Aust. 

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1918. A check list of the marine fauna of New South Wales. 1. Suppl. to the /. roy. 

Soc. N.S.W. 51 : 1-114. 

1923. Studies on Australian Mollusca. Part 14. Proc. linn. Soc. N.S.W. 48 : 301-316. 

Heilprin, A. 1886. A new species of Aplysia. Proc. Acad. nat. Sci. Philad. Part 3 : 364. 

1888. Contributions to the Natural History of the Bermuda Islands. Ibid. : 302-328. 

Hertling, H. 1935. Der Seehaue (Aplysia) bei Helgoland. Natur u. Volk. 65, 3 : 103-108. 
Hoffmann, H. 1926. Opisthobranchia in Die Tierwelt der Nord- und Ostsee, 3, 9 : 1-66. 

1939- Opisthobranchia in Bronn's Klassen und Ordnungen des Tier-reichs. 3 [2(3)] : 1-1247. 

Howells, H. H. 1942. The structure and function of the alimentary canal of Aplysia 

punctata. Quart. J. micr. Sci. 83 : 357-397. 
Hunt, A. R. 1877. On some large Aplysiae taken in Torbay in 1875. Trans. Devonsh. Ass. 
9 : 400-403. 

1878. On the growth of Aplysiae in Torbay. Ibid. 10 : 611-617. 

Hutton, F. W. 1875. Description of two new species of Aplysia. Trans. Proc. N.Z. Inst. 7 : 279. 

1880. Manual of the New Zealand Mollusca. Wellington: 1-224. 

1882. Notes on some branchiate Gastropoda. Trans. N.Z. Inst. 15 : 1 18-130. 

1904. Index Faunae Novae Zealandiae. London, for Phil. Inst. Canterbury, N.Z. : 

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International Commission on Zoological Nomenclature. 1954. Opinion 200. Valida- 
tion under the plenary powers, of the accustomed usage of the generic names Tethys L. 1767 
and Aplysia L. 1767. Opin. int. Comm. zool. Nom. 3 : 239-266. (Aplysia L. 1767 is 
placed on the Official List of genera with type Aplysia depilans Gmelin, 1791. Usage of 
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1955a. Opinion 354. Addition to the Official List ... of the specific name " fasciata " 

Poiret, 1789, as published in the combination "Laplysia [recte " Aplysia "] fasciata [Class 
Gastropoda]. Opin. int. Comm. zool. Nom. 11 : 81-90. 

1 955°- Opinion 355. Validation, under the plenary powers, of the specific name punctata 

Cuvier, 1803, as published in the combination Laplysia (recte Aplysia) punctata (Class 
Gastropoda). Opin. int. Comm. zool. Nom. 11 : 91-102. 



REVISION OF THE WORLD SPECIES OF APLYSIA 397 

International Commission on Zoological Nomenclature. 1959. Opinion 560. Protection 
under the Plenary Powers of the specific name parvula Morch, 1863, as published in the com- 
bination Aplysia parvula (Class Gastropoda). Opin. int. Comm. zool. Nom. 20 : 293-302. 

Jeffreys, Gwyn. 1866. Report on dredging among the Channel Islands. Rep. Brit. Ass. 
Birmingham, 1865, 35, Rep. on the state of Sci. 1-3. 

1869. British Conchology , 5 : 1-259. London. 

Jhering, H. von. 191 5. Die Opisthobranchien der brasilianischen Kiiste. Nachr. Bl. 

dtsch. Malako-zool. Ges. 47 : 133-144. 
Johnston, G. 1849. A descriptive catalogue of the Gasteropodous Mollusca of Berwick- 
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Jordan, H. 1917. Das Wahrnehmen der Nahrung bei Aplysia limacina und Aplysia depilans. 

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Jullien, A. 1936. Automatisme des lambeaux cardiaques chez Aplysia fasciata. C. R. Soc. 

biol. Paris, 123 : 905-906. 
Kirk, T. W. 1882. Description of new Shells. Trans. Proc. N.Z. Inst. 14 : 282-283. 
Krauss, F. 1848. Die sudafrikanischen Mollusken. Stuttgart : 1-72. 
Lamarck, J. B. P. A. 1822. Histoire naturelle des Animaux sans Vertebres. 6, part 2. 

Aplysiens, 37-40. 
Lankester, E. R. 1875. Contributions to the developmental history of the Mollusca. 2. The 

early development of two species of Aplysia. Phil. Trans, roy. Soc. 165 : 13-27. 
Leach, W. E. 1847. The classification of the British Mollusca. Ann. Mag. nat. Hist. 20, 

2 : 267-273. 

1852. A Synopsis of the Mollusca of Great Britain. London : 1-376. 

Lesson, R. P. 1830. Zoology in L. I. Duperrey Voyage autour du Monde, Corvette La Coquille. 

2 : 294-5. 
Letson, E. 1898. Description of a new Tethys [Aplysia) pilsbryi. Proc. Acad. nat. Sci. 

Philad. : 193, pi. viii. 
Lo Bianco, S. 1888. Notizie biologiche riguardanti specialmente il periodo di maturita 

sessuale degli animali del Golfo di Napoli. Leipzig. Mitt. zool. Sta. Neapel. 8 : heft 3 

and 4, 385-440. 
Locard, A. 1886. Prodrome de Malacologie francaise. Mollusques marins. Ann. Soc. 

Agric. Lyon. 5th ser. 8 : 1-216. 
Lonnberg, E. 1933. Zur Kenntnis der Carotinoide bei marinen Evertebraten. Ark. Zool. 

25 : 1-17. 
Lowe, H. N. 1934. O n tne Sonoran side of the Gulf. Nautilus, 48, 2 : 1-4 and 43-46. 
MacFarland, F. M. 1909. The Opisthobranchiate Mollusca of the Branner-Agassiz Expedi- 
tion to Brazil. Leland Stanf. Univ. Publ. Univ. Ser. 2, 5-104. 

1924. Opisthobranchiate Mollusca. Expedition of the Californian Academy of Science 

to the Gulf of California in 192 1. Proc. Calif. Acad. Sci. 4, 13, No. 25 : 389-420. 
Macginitie, G. E. 1934. The egg-laying activities of the sea hare, Tethys californicus 

(Cooper). Biol. Bull., Woods Hole, 67 : 300-303. 
Macnae, W. 1955. On four species of the genus Aplysia common in South Africa. Ann. 

Natal. Mus. 13 : 223-241. 

1957- O n the occurrence of Aplysia dactylomela and Aplysia oculifera in South Africa. 

5. Afr. J. Sci. 53, 11 : 289-292. 

Marcus, E. 1956. Notes on Opisthobranchia. Bol. Inst. Oceanogr., Sao Paulo, 7 : 31-63. 

1958. On western Atlantic Opisthobranchiate Gastropods. Amer. Mus. Novit. No. 

1906 : 1-82. 

Marcus, E. & E. 1955. Sea hares and side-gilled slugs from Brazil. Bol. Inst, oceanogr. 
Univ. Sao Paulo, 6 : 3-48. 

1959- Notes on Aplysia. Ibid. 8, 3-20. 

Martens, E. von. 1880. Mollusken in K. MQbius : Beitrdge zur Meeresfauna der Insel 
Mauritius und der Seychellen. Berlin : Mollusca : 181-352 ; taf. 19-22. 

1 894-1 903. Mollusken in R. Semon : Zoologische Forschungen Reisen in Australien 

und der Malay Archipel. Denskschr. med, naturw, Ges. Jena, 8 : 83-96. 



398 REVISION OF THE WORLD SPECIES OF APLYSIA 

Mazzarelli, G. 1889. Intorno alle secrezioni della glandola opalina (Vayssiere) e delle 
glandola dell opercolo branchiale nelle Aplisie del Golfo di Napoli. Zool. Anz. 
12 : 580-583. 

1890. Nachricht iiber eine neue Art von Aplysia {A. lobiancoi). Nachr. Bl. dtsch. malako- 

zool. Ges. 22 : Nos. 3 and 4, 42-43. 

1 89 1. Ricerche sulla morfologia e flsiologia dell' apparato riproduttore nelle Aplysiae 

del Golfo di Napoli. Atti. Accad. Sci. fis. nat. Napoli, 4, ser. 2 : 1-50. 

1893. Monografia delle Aplysiidae del Golfo di Napoli. Mem. Soc. ital. Sci. nat. (3), 

9 : 1-222. 

Mazzarelli, G. & Zuccardi, R. 1889. Su di alcune Aplysiidae dell 'Oceano Pacifico, apparte- 

nenti alia collezione Chierchia. Boll. Soc. nat. Napoli, 3 : 47-54. 
Mazzarelli, G. & Zuccardi, R. 1892. Sulle Aplysiidae raccolte dal tenente di vascello G. 

Chierchia nel viaggio della V. Pisani, 1882-85. Mem. Soc. ital Sci. nat. (3), 8 : 1-21. 
Meinertzhagen, F. H. 1880. Note and description of a possibly new species of Aplysia (tryonii). 

Trans, and Proc. N.Z. Inst. 12 : 270-271. 
Merriman, D. 1937. The occurrence of Tethys willcoxi in New England waters and another 

record for this locality. Nautilus, 50 : 95-97. 
Mobius, K. A. 1880. See Martens, E. von. 
Morch, O. A. L. 1863. Contributions a la Faune malacologique des Antilles danoises. /. 

Conchy liol. y s6rie, t. 3", 11 : 21-43. 

1875. Synopsis Molluscorum marinorum indiarum occidentalium. Malakozool. Bl. fur 

1874 and 5, 22 : 142-202. 

Norman, A. M. 1890. Revision of British Mollusca. Ann. Mag. nat. Hist. (6), 6 : 60-91. 
Odhner, N. H. 1907. Northern and Arctic Invertebrates. 3. Opisthobranchia and Pteropoda. 
K. Svensk. vetensk. Akad. Handl. 41, 4 : 1-118. 

1922. Norwegian Opisthobranchiate Mollusca in the collections of the Zoological Museum 

of Kristiania. Medd. zool. Mus. Kristiania, No. 1 : 1-47. 

1932. Beitrage zur Malakozoologie der Kanarischen Inseln. Ark. Zool. 23 A, No. 

14 : 1-116. 
O'Donoghue, C. H. 1924. Report on the Opisthobranchiata from the Abrolhos Islands, 

Western Australia. /. Linn. Soc. (Zool.) 35 : 521-579. 

1929. Opisthobranchiate Mollusca collected by the South African Marine Biological 

Survey. Rep. Fish. mar. biol. Survey S. Afr. 7 : 1-84. 

Olmsted, J. M. D. 1918. Notes on the locomotion of certain Bermudian Molluscs. /. exp. 

Zool. 24 : 223-236. 
d'Orbigny, A. 1839 (text) and 1836-44 (pis.). Mollusques in Webb & Berthelot's Histoire 
Naturelle des lies Canaries 2 (2) : 3-1 17, pis. 1-7 and 7B. 

1837. Voyage dans V Amerique Meridionale, 1826-33, 5 : 1-758. 1847. Atlas, 9 : Pis. 1-85. 

— — 1854. Histoire ... de Vile de Cuba (Ramon de la Sagra) Paris, 1853, 1 : 117. List of 

the shells of Cuba in the collection of the British Museum, London. 
Ostergaard, J. M. 1950. Spawning and development of some Hawaiian marine gastropods. 
Pacif. Sci. 4 : 75-115. 

1955- Some Opisthobranchiate Mollusca from Hawaii. Ibid. 9 : 1 10-136. 

Parker, G. H. 1918. The pedal locomotion of the Sea Hare, Aplysia californica. J. exp. Zool. 

24, 1917-18 : 139-145- 
Payraudeau, B. C. 1826. Catalogue descriptif et methodique des Annelides et des Mollusques 

de Vile de Corse. Paris. 1-2 18. 
Pease, W. Harper, i860. Description of new species of Mollusca from the Sandwich Islands. 

Proc. zool. Soc. Lond. 28 : 18-36. 

1868. Descriptions of marine Gastropodae inhabiting Polynesia. Amer. J. Conch. 4 : 71-80. 

Pennant, T. 1812. British Zoology. 5th Edition. London, 4 : 1-379. 

Perrier, R. & Fischer, H. 1908. Anatomie et histologic comparees des glandes de Blochmann 

chez les Tectibranches. C.R. Acad. Sci. Paris, 146 : 1335-1337. 
Philippi, R. A. 1836 and 1844. Enumeratio molluscorum siciliae. Berlin. (Vol. 2, Halis Saxonum, 

1844 : 98, species of Aplysia.) 



REVISION OF THE WORLD SPECIES OF APLYSIA 399 

Pilsbry, H. A. 1895. O n the status of the names Aply'sia and Tethys. Proc. Acad. nat. Sci. 

Philad. : 347. 

1895-96. Manual Conch. 16, part 62, 1895 : 1-112 ; part 63, 1896 : 113-262. 

1951. Aplysia badistes, a peculiar floridean sea-hare. Notul. Nat. Acad. Philad. 240 : 1-6. 

Pliny, C, a.d. 60. Historia naturalis. Lib. 9, 32. 

Poiret, J. L. M. 1789. Voyage en Barbaric Paris, 2 : 1-3 15 (2 e pt.). 

Pruvot-Fol, A. 193 i. Notes de systematique sur les Opisthobranches. Le statut d' Aplysia 

et de Tethys. Bull. Mus. Hist. nat. Paris (2) 3 : 308-316. 

1932. Du genre Aplysiopsis Bergh. Ibid. (2), 4 : 322-323. 

i933#- Les Opisthobranches de Quoy et Gaimard. (Note pr61iminaire.) Ibid. 5 : 400-401. 

19336- Note sur les tentacules des Aplysiad6s et sur la systematique de ce groupe. Bull. 

Soc. zool. Fr. 58 : 99-105. 
1934- Les Opisthobranches de Quoy et Gaimard. Arch. Mus. Hist, nat Paris (6). 11 : 13-92. 

1935- De quelques suppressions de genres jug6s inutile, parmi les Mollusques Opistho- 
branches. Note de syst6matique, No. 14. Bull. Mus. Hist. nat. Paris (2), 7 : 254-257. 

1947- Les Opisthobranches de W. Harper Pease. Revision. /. Conchyliol. 87 : 96-114. 

1949. Une derniere note au sujet du nom de Tethys. Ibid. 89 : 94. 

1953. Etude de quelques opisthobranches de la cote atlantique du Maroc et du S6n6gal. 

Trav. Inst. sci. Gherifien, No. 5 : 3-105. 

1954- Etude d'une petite collection d'opisthobranches d'oc6anie francaise. /. Conchyliol. 

94 : 3-30. 

Quoy, J. R. C. & Gaimard, P. 1832. Zoology, 2, in Voyage de decouvertes de V Astrolabe, 1826-9. 

Paris : 308-316. Atlas, 1833. 
Rang, Sander. 1828. Histoire naturelle des Aplysiens, premiere famille de l'ordre des Tecti- 

branches. In Ferussac : Hist. nat. gen. etpartic. des Mollusques. Paris 1827 : 1-84, 24 plates. 
Rathke, J. 1799. Jagttagelser henhorende til Involdeormene og Bloddyrenes Natur- 

historie. Skriv. nat. Hist. Selsk., Copenhagen, 5, heft. 1 : 61-148. 
Redi, F. 1684. Opere 2. Osservazioni di Francesco Redi . . . intorno agli animali viventi 

che si trovano negli animali viventi. Firenze. 243 pp. 
Regan, C. T. 1913. The fishes described by Carmichael from Tristan da Cunha. Ann. Mag. 

nat. Hist. (8), 11 : 466-468. 
Risbec, J. 1928. Etude anatomique des Gasteropodes Tectibranches de la presqu'ile de Noumea. 

Arch. Mus. Hist. nat. Paris (6), 3 : 37-68. 

1929. Notes zoologiques et anatomiques sur quelques Opisthobranches de Madagascar. 

Faune Colon, franc. 3 : 45-62. 

1 93 1. Etude de quelques gasteropodes opisthobranches de la cote atlantique du Maroc. 

Bull. Soc. Sci. nat. Maroc, 11 : 67-89. 

1 95 1. Notes sur les Tectibranches de Nouvelle Caledonie. /. Soc. Oceanist. 7 : 123-158. 

Risso, A. 1818. Memoire sur quelques gasteropodes nouveaux nudibranches et tectibranches 
observes dans la mer de Nice. /. phys. chim. hist. nat. et des arts, 87 : 368-377. 

1826. Histoire naturelle des principales productions de I'Europe meridionals. Mollusques. 

Paris, 4 : 1-439. 

Rizzi, M. 1908. Sullo svillupo della radula nel genere Aplysia. Atti. 1st. Veneto, 68 : 2, 261-274. 
Rochebrune, A. T. de. 1881. Mat6riaux pour la faune d'Archipel du Cap Vert. Nouv. Arch. 

Mus. Hist. nat. Paris (2), 4 : 215-340, pi. 17-19. 
Rondelet, G. 1554. D e Piscibus Marinis. Lugdini : 583. Aplysia fasciata as Lepus marinus : 

520-526. 
Ruppell, E. & Leuckart, F. S. 1828. Neue wirbellose Thiere des rothen Meers. Part 5 in 

Ruppell E. Atlas zu der Reise im nordlichen Afrika. Senckenberg. Naturf. Ges. Frankfurt- 

am-Main : 1-47, pis. 1-12. 
Saint Loup, R. 1888. Notes et observations anatomiques sur les Aplysies. C.R. Acad. Sci. 

Paris, 107 : 1010. 
Sanford, S. N. F. 1922. Tethys willcoxi in New England waters. Nautilus, 35 : 80-82. 
Sars, G. O. 1878. Bidrag til Kundskaben om Norges Arktiscke Fauna. 1. Mollusca. Regionis 

Arcticae Norvegiae. Christiania : 1-466. (Figure of A. punctata only, Tabl 12.) 



4 oo REVISION OF THE WORLD SPECIES OF APLYSIA 

Sars, M. 1840. Beitrag zur Entwickelungsgeschichte der Mollusken und Zoophyten. Arch. 

Naturgesch. 6 : 196-219. 
Sinel, J. 1906. An outline of the Natural History of our Shores. London: 1-123. 
Smith, E. A. 1884. Report on the zoological collections made in the Indo-Pacific ocean during 

the voyage of H. M.S. Alert, 1881-82. London : i-xxv, 1-684, pis. 1-54. Mollusca : 34-116, 

pis. 1-7. 

1903. A list of species of Mollusca from South Africa, forming an appendix to G. B. 

Sowerby 's Marine Shells of South Africa. Proc. malac. Soc. Lond. 5, No. 6 : 354-402, pi. 15. 

Sowerby, G. B. 1869. Aplysia in Reeve A. L. Conchologia Iconica, 17 : plates only : 1-10. 
Sowerby, James. 1804-06. British Miscellany. 2 vols, in 1. 4 . London. R. Taylor & Co. 

76 col. pi. 
Stimpson, W. 1855. Descriptions of some of the new marine Invertebrata from the Chinese 

and Japanese Seas. Proc. Acad. nat. Sci. Philad. 7 : 375-380. 
Summer, F. B., Osburn, R. C. & Cole, L. J. 191 1. A biological survey of the waters of Woods 

Hole and vicinity. Part 2, Sect. 3, A catalogue of the marine fauna. Bull. U.S. Bur. Fish. 

31 : 1-794- 
Suter, H. 1913 and 1915 (Plates). Manual of the New Zealand Mollusca. Wellington, N.Z. 

1-1120. PI. 1-72. 
Swainson, W. 1840. A Treatise on Malacology. London, Longmans : 1-419. 
Sykes, E. R. 1905. The Molluscs and Brachiopods of Bally nakill and Bonn Harbours, 

County Galway. The marine fauna of the west coast of Ireland, Part 2. Rep. Sea Inl. 

Fish. Ire. Sci. Invest, for 1902-03 : 53-92. 
Takahashi, S. 1934. Ecological notes on Tethys {Aplysia) punctata in Formosa. Venus, 

4 : 357-36o. 
Taki, Iw. 1932. On Tethys dactylomela. Ibid. 3 : 212 and 351. 
Tarao, Sh. 1934. Microscopical study on the purple gland of Tethys punctata Cuv. /. Fac. 

Sci. Hokkaido Imp. Univ. (6), Zool. 3 : 221-231. 
Tchang, Si, 193 1. Contribution a V etude des Mollusques Opisthobranches de la cote provencale. 

These. Lyon : 1-22 1. 
Tenison Woods, J. E. 1876. Description of new Tasmanian Shells. Rep. Roy. Soc. Tas- 
mania for 1875 : 134-162. 
Thiele, J. 1910. Molluskenfauna Westindiens. Zool. Jb. Suppl. 11 : 109-132. 

1925. Gastropoda der deutschen Tiefsee Expedition. 2. Wiss. Ergebn. " Valdivia ", 

17 : 36-382. 

1929. Handbuch der systematischen Weichtierkunde. Jena. 2 : 377-778. 

1930. Gastropoda und Bivalvia in Die Fauna Sud-west Australiens, 5, 8 : 561-596. 

Michaelsen und Hartmeyer. 

Thompson, W. 1845. Additions to the fauna of Ireland. Ann. Mag. nat. Hist. 15 : 308-321. 

1856. Natural History of Ireland. London. 4 : 1-5 16. 

Tilesius von Tilenau, W. G. 1813. In A . J. von Krusenstern's Reise um die Welt in den 
Jahren, 1803-06. [Not seen, the copies in London, Oxford and Paris are incomplete. 
Rang quoted the text and copied the figure.] 

Tryon, G. W. Manual of Conchology. See Pilsbry, H. A. 

Turton, W. H. 1932. The marine shells of Port Alfred, South Africa. Oxford and London : 
i-xvi and 1-33 1. 72 pis. 

Verany, G. B. 1853. Catalogue des Mollusques, Cephalopodes, Pteropodes, Gasteropodes, 
Nudibranches etc. . . . des environs de Nice. /. Conchy Hoi. 4 : 375-392. 

Vayssiere, A. 1885. Recherches zoologiques et anatomiques sur les Mollusques opistho- 
branches du Golfe de Marseille. 2, Tectibranches. Ann. Mus. Hist. nat. Marseille : 5-18 1. 

1935- Faune Flore Mediterraneenne. Paris, No. 20. 

Verrill, A. E. 1900. The Nudibranchs and naked Tectibranchs of the Bermudas. Trans. 
Conn. Acad. Arts Sci. for the years 1899-1900, 10, pt. 2 : 545-550, pi. LXVI. 

1 901. Additions to the Fauna of the Bermudas, from the Yale Expedition of 1901, with 

notes on other species. Ibid. 11 : 15-62. 

Webb, P. B. & Berthelot, S. 1835-44. See d'Orbigny, A. 



REVISION OF THE WORLD SPECIES OF APLYSIA 



401 



White, K. M. 1952. On a collection of molluscs from Dry Tortugas, Florida. Proc. 
malac. Soc. Lond. 29 : 106-120. 

1955- Some Opisthobranchs from West Africa. Res. Scient. Exped. Oceanogr. Belg. 

Eaux Cot. Afr. 3 (4) : 163-195. 

Winckworth, R. 1927. The littoral fauna of Krusadai Island in the Gulf of Manaar. 

Mollusca. Bull. Madras Govt. Mus. : 95-109. 
Winkler, L. R. 1955. A new species of Aplysia on the southern California coast. Bull. S. 

Calif. Acad. Sci. 54 : 5-7. 
1958. Metamorphosis of the shell in the California sea hare, Aplysia califomica Cooper, 

Pacific Sci. 12 : 348-349. 

1959a. A mechanism of color variation operating in the west coast sea hare, Aplysia 

califomica Cooper. Pacific Sci. 13 : 63-66. 

1959b. A new species of sea hare from Californian waters. Bull. S. Calif. Acad. Sci. 

58 (1) : 8-10, pi. 5. 



oculifera 



INDEX TO SPECIFIC NAMES 



adamsi Pilsbry, 1896 (unidentifiable) .... 

alba Cuvier, 1803 { — fasciata) ..... 

albo -punctata Deshayes, 1853 (= punctata) . 

allochroa Bergh, 1908 non O'Donoghue, 1929 (= parvula) 

allochroa ; O'Donoghue, 1929 as Tethys, non Bergh, 1908 ( 

angasi Sowerby, 1869 (= dactylomela) 

anguilla Cuming in Sowerby, 1869 ( = parvula) 

annulifera Thiele, 1930 (= dactylomela) 

aperta Turton, 1932, as Tethys (= dactylomela) 

argus Ruppell & Leuckart, 1828 (= dactylomela) 

atromarginata Bergh, 1905 (= parvula) 

australiana Clessin, 1899 (= parvula) 

badistes Pilsbry, 195 1, as Tethys ( = Juliana) 

benedicti Eliot, 1899 (= dactylomela) . 

bipartita Turton, 1932, as Tethys (unidentifiable) 

bipes Pease, i860, as Syphonota ( = Juliana) 

brasiliana Rang, 1828 .... 

brunnea Hutton, 1875 (= nigra var. brunnea) 

cailleti Deshayes, 1857 (= ? brasiliana) 

califomica Cooper, 1863 .... 

cameliformis Locard, 1886 ( = fasciata) 

camelus Cuvier, 1803 (= fasciata) 

capensis O'Donoghue, 1929 as Tethys ( = Juliana 

cedrosensis Bartsch & Rehder, 1939 . 

cervina Dall & Simpson, 1902 . 

chierchiana Mazzarelli & Zuccardi, 1892 (unidentifiable) 

concava Sowerby, 1869 (= parvula) . 

comigera Sowerby, 1869 . 

cronullae, nov. .... 

cuvieri delle Chiaje, 1822 (= punctata) 

cuvieriana delle Chiaje, 1822 ( = 

dactylomela Rang, 1828 . . 271-2, 274 

d. var. aequorea Heilprin, 1888 . 

d. var. fimbriata Adams & Reeve, 1850 

delli, nov. (= nigra var. delli) . 

denisoni Smith, 1884 .... 



276-9, 289, 307 



75- 



10, 3 



Page 

• 273 
315-6, 318 

291 
. 287 

• 332 
307, 309 
287, 291 

307 
307 
307 
287 

287 

363 

307, 309-10 

. 274 

-365» 367-9 
342-3, 382 

369-7° 

. 297 

283, 294-7 

• 315 
315-6, 318 

• 363 
277. 354-6, 363, 374 

272, 276, 299-302, 382 
273, 277 
287, 291 
272, 281, 302-4, 337 
. 272, 304-6, 347 
291 
291 
2 4> 332-3, 335 337. 382 

• 307 
. 308 

• 370 
272, 310-12 



277, 

363 
297-300, 302 



71. 277 



4-02 



REVISION OF THE WORLD SPECIES OF APLYSIA 



271, 274-5, 277, 292, 294 



1 deperdita Philippi, 1844 (not an Aplysia) 

depilans Gmelin, 1791 

depilans ; Pennant, 1812 {—punctata) . 

depilans; de Blainville, 1823 ( = fasciata) . 

depilans major Barbut, 1783, as Laplysia (= depilans) 

depilans minor Barbut, 1783 (= punctata) . 

dumortieri Cantraine, 1840 (= punctata juv.) 

dura, nov. ....... 

elegans Turton, 1932 (unidentifiable) . 

elongata Pease, i860 as Siphonota (unidentifiable) 

euchlora A. Adams, 1861, and euchlorus Adams in Gray, 185 

euclorus as Aclesia ; M. E. Gray, 1850 (= euchlora) 

eusiphonata Bergh, 1908 (= maculata) 

excavata Sowerby, 1869 (= Sydney ensis) 

extraordinaria Allan, 1932 

fasciata Poiret, 1789 .... 

ferussaci Rang, 1828 (= punctata) 

fimbriata Adams & Reeve, 1850 (= dactylomela) 

floridensis Pilsbry, 1895, as Tethys (= brasiliana) 

fragilis Lamarck, 1822, as Dolabella (= depilans) 

fusca Tilesius, 181 2 (unidentifiable) 

gargantua Bergh, 1908 ( = maculata) 

gigantea Sowerby, 1869 

gilchristi Bergh, 1908 (= maculata) 

gracilis, nov. .... 

grandis Pease, i860, as Siphonata (unidentifiable) 

griffithsiae Gray, 1850 (= punctata juv.) 

griffithsiana Leach, 1852, as Esmia (= ? punctata juv.) 

guadeloup ensis Sowerby, 1869 (= brasiliana or dactylomela) 

guttata M. Sars, 1840 (= punctata) 

hamiltoni Kirk, 1882 ( = nigra) 

hirasei Baba, 1936, as Tethys ( = oculifera) 

hyalina Sowerby, 1869 (= nigra) 

hybrida J. Sowerby, 1806 (= punctata) 

immunda Bergh, 1902 (= euchlora) 

inca d'Orbigny, 1837 

intermedia Farran, 1905 (= parvula) 

japonica Sowerby, 1869 (= parvula) 

juanina Bergh, 1898, as Aplysiopsis (= parvula) 

Juliana Quoy et Gaimard, 1832 . . . 271, 

Juliana var. bipes 

j. var. quoy ana 

j. var. sibogae .... 

julienna mis-spelling of Juliana, as in H. and A. Adams 

keraudreni Angas, 1867, as Syphonota non Rang, 1828 (= dactylomela) 

keraudveni Rang, 1828 ..... 

kurodai Baba, 1937 • 

laevigata Stimpson, 1855 (unidentifiable) 

lavis de Blainville, 1823, as Dolabella (— depilans) 

leporina Rang, 1828 ( = fasciata) 

leporina delle Chiaje, 1823, as Aplisia (= depilans) 

lepus Risso, 1826, as Dolabella ( = fasciata) 

1 This is a fossil shell, which is not an Aplysia. Depontaillier (i 
the linings of the shells of Pectunculus and Lucina. 



277, 



Page 

• 395 
316, 357-6o, 365 

291 
315 
357 
291 
291 
356, 360-3 

• 274 
273, 277, 287-8, 291 

271-2, 374-7 

• 374 
. 326 

• 348, 350 

2 75-6, 312 

272, 275, 315-8, 357, 382 

292 

307 
297 

357 

273 

326 

272, 318-20 

. 326 

320-1 

273. 307 
292 

274, 292 

297, 307 

292 

369-70 

332, 334 
369-70 
. 292 

• 374 
272, 277, 321-3 

287 

287 

287 

290, 356, 359, 363-9, 37° 

363, 365, 368 

363, 368 

363, 368 

• 368 

307, 324 

273, 277, 281, 322-5 

272, 325-6 

• 273 

• 357 
315-8 

• 357 

• 315 
says Philippi' s specimens are only 



S3, 



REVISION OF THE WORLD SPECIES OF APLYSIA 



4°3 



lessoni Rang, 1828 (= ? keraudreni) . 

limacina Blainville, 18236, (= fasciata) 

limacina L. 1758 as Tethys (= depilans) 

lineolata Adams & Reeve, 1850 ( = ? oculifera 

livida d'Orbigny, 1837 (= brasiliana) 

lobata Bergh, 1908 (= dactylomela or parvula juv 

lobiancoi Mazzarelli, 1890, as Siphonota ( = fasciata or a 

species) .... 
longicornis Rang, 1828 (= punctata) 
lurida mis-spelling of livida 
macula Turton, 1932 (unidentifiable) 
maculata Rang, 1828 
major Lankester, 1875 (= depilans) 
marginata A. Adams, 1861 (unidentifiable) 
marginata de Blainville, 1823 (= punctata) 
marinus Rondelet, 1554, as Lepus ( = fasciata) 
marmorata de Blainville, 1823 ( = fasciata) 
marmorea A. Adams, 1861 (unidentifiable) . 
megaptera Verrill, 1901 (= dactylomela) 
melanopus Couch, 1870 (= depilans) . 
modesta Thiele, 1910, as Tethys (= ? morio juv 
monochroa Bergh, 1908 (= parvula) . 
morio Verrill, 1901 . 
mouhoti Gilchrist, 1895 (not an Aplysia) 
mustelina Davies in Pennant, 1812 (= ? punctata) 
napolitana delle Chiaje, 1823 ( = fasciata) . 
neapolitana delle Chiaje, 1823 ( — fasciata) . 
nettiae Winkler, 1959 (= ? calif ornica) 
nexa Thompson, 1845 (= punctata) 
nigra d'Orbigny, 1837 .... 
nigrocincta Martens, 1880 (= parvula var.) 
nigromarginata Risso, 18 18 (= punctata) 
nodifera Adams & Reeve, 1850 (= oculifera 
norfolkensis Sowerby, 1869 (= parvula) 
ocellata A. Adams, 1861 non d'Orbigny, 1837 (unidentifiable) 
ocellata d'Orbigny, 1837, non A. Adams, 1861 ( = dactylomela) 
oculifera Adams & Reeve, 1850 
odorata Risbec, 1928 ( = dactylomela) 
odorata var. bourailli Risbec, 1951 (= dactylomela) 
operta Burne, 1906 (= dactylomela) . 
orientalis Sowerby, 1869 (= parvula) 
paeteliana Clessin, 1899 ( = parvula) . 
panamensis Pilsbry, 1895, as Tethys (— dactylomela) 
parva Pruvot-Fol, 1953 (juvenile, = Juliana) 
parvula (Guilding) Morch, 1863 • • 271-2 

peasei Pilsbry, 1895, as Tethys (= parvula) 
peregrina Thiele, 1925 (= parvula) 
peter sonii Gray, 1828 (= depilans) 
petiti Risbec, 1929 (= ? Juliana juv.) 
pilsbryi Letson, 1898 (= cervina) 
piperata Smith, 1884 (not an Aplysia) 
poikilia Bergh, 1908 (= maculata) 
poli delle Chiaje, 1823 (= depilans) . 
poliana delle Chiaje, 1823 (= depilans) 



hybrid between two 



276- 



7> 283 



, 287 



Page 

3 22 , 3 2 5 

315, 318 

. 274 

• 332 
297, 299 
287, 307 

Mediterranean 

274, 294, 315 

292 

297, 299 

. 274 

275, 287-8, 326-8 

• 357 

• 273 
292, 294 

• 274, 315 

• 315 

• 273 
307, 310 

• 357 
. 328 

. 287 

275-6, 328-32 

■ 374. 39i 
. 292 

315-6 

• 315 
. 294 
. 292 

277» 369-7° 

277-8, 287, 291 

292 

332, 334 
287-8, 291 

307 

307 

272, 279, 281, 332-4 

307 
307 
307 
287 
287 
307 
273 

291. 327, 343. 345- 38o 
287, 291 
. 287 

• 357 

• 363 
. 299 

• 39i 

326, 328 

• 357 

• 357 



4°4 



REVISION OF THE WORLD SPECIES OF APLYSIA 



protect Rang, 1828 = (dactylomela) 

pulmonica Gould, 1852 

pulmonica var. tryoniana Pilsbry, 1895, as Tethys 

punctata Cuvier, 1803 ..... 

punctata; Takahashi, 1934 (probably k urodai) . 

punctata Pease, i860, as Syphonota (may be parvula) 

punctatella Bergh, 1902 (unidentifiable) 

quoyana Engels & Eales, 1957 (var. of Juliana) . 

radiata Crouch, 1826 ( = fasciata) 

radiata Ehrenberg, 1831 (= dactylomela) . . 

rangiana d'Orbigny, 1837 (= Juliana juv.) . 

rehderi, nov. ....... 

reticulata, nov. ...... 

ritteri Cockerell, 1902 (= californica) 

robertsi Pilsbry, 1895, as Tethys 

rosea Rathke, 1799 (= punctata juv.) 

rosea; Sowerby, 1869 non Rathke (= parvula) 

sagamiana Baba, 1949 ..... 

sandvichensis Sowerby, 1869 ( = Juliana) 

schrammii Deshayes, 1857 (= dactylomela) . 

scutellata Ehrenberg, 1831 (= dactylomela) . 

sibogae Bergh, 1905 ( = Juliana var.) . 

sicula Swainson, 1840 (= ? fasciata) . 

sieboldiana Clessin, 1899 (= parvula) 

sinensis Sowerby, 1869 (unidentifiable) 

sorex Rang, 1828 (juvenile of doubtful status) 

sowerbyi Pilsbry, 1895, as Tethys (replaces Sowerby 's tigrina 

sparsinotata Smith, 1884 (not an Aplysia) 

spuria Krauss, 1848 (unidentifiable) . 

stellata Risso, 1818 (= punctata) 

subquadrata Gould, in J. Sowerby, 1806 (= punctata) 

subquadrata ; Sowerby, 1869, non Gould (= parvula) 

sydney ensis Sowerby, 1869 .... 

tarda Verrill, 1901 (unidentifiable) 

tasmanica Tenison Woods, 1876 (= gigantea) 

tigrina Rang, 1828 (= dactylomela) 

tigrina Quoy et Gaimard, 1832, non Rang, 1828 (= tigrinella 

tigrina Angas, 1867, non Rang, 1828 (= sowerbyi) 

tigrinella M. E. Gray, 1850, new name for A. tigrina Q 

trigona Sowerby, 1869 (= parvula) 

tryonii Meinertzhagen, 1880 (unidentifiable) 

unicolor Risso, 181 8 (= punctata) 

vaccaria Winkler, 1955 

varians Leach, 1852 (= punctata) 

velifer Bergh, 1905 (= ? dactylomela juv.) 

vensoa Hutton, 1875 (unidentifiable) . 

vexans Bergh, 1905 (unidentifiable) . 

virescens Risso, 1826 (= punctata) 

viridescens Pease, 1868, as Siphonota (= dactylomela) 

vistosa Pruvot-Fol, 1953 (unidentifiable juvenile) 

vulgaris de Blainville, 1823 (= fasciata) 

willcoxi Heilprin, 1886 

w. var. perviridis Pilsbry, 1895, as Tethys 

winneba Eales, 1957 

woodii Bergh, 1908 ( = Juliana) 



Page 

307» 3°9-io 

272, 277, 323, 335-7, 382 

335 
1, 275-6, 279, 284, 288, 291-5, 325, 380 

325 

287 

273 
368 

3i5 

307 

277> 363, 369 

272, 337-4° 

340-1, 348 

. 294 

342-3 
279, 292, 294 

. 287 
272, 343-5 



et G 



non Rang, non Q 



Gray 

,183 



72, 306, 340, 345 



1850 




382 

• 363 

• 307 

• 307 
. 363. 368 

• 315 

. 287 

• 273 
274. 363. 369 

etG.) 272,345-7 

• 39i 
273, 287, 291 

. 292 
. 292 
. 287 

347. 348-50. 364 

• 273 
. 318 

307, 309-10 
maculata) 288, 326 

345 

(= maculata) 288, 326, 328 
287, 291 
273, 276, 287, 291 
. 292 
• 277, 371-4 
292 

• 307 

• 273 

• 273 
. 292 

• 307 
. 274 

• 3i5 
272, 276, 350-2 

. 350. 352 
. 272, 352-4 
. 363. 382 



INDEX TO VOL: V 



The page numbers of the principal references and the new taxonomic names are printed in 

Clarendon type. 



abstrusus, Prosciurillus . 
aelocephalus, Haplochromis 

africana, Physopsis 

ai, Sagitta 

alfari, Microsciurus 

alluaudi, Astatoreochromis 



65, 167 



264 
214-217 
8, 19, 22 
232-233 

263 
■173, 188 



alluaudi, Astatoreochromis alluauda 



173, 



174-177 

• 43 

55-70 

267-404; 270, 272, 380-382 



Anaspidacea 
Anhinga 
Aplysia (genus) 
Aplysia (sub-genus) 

271, 272, 273, 279, 354-374. 3§2 
Aplysiidae . . . . . .269 

Aplysiinae ...... 269 

arctica, Nidalia . . . . .112 

Ardeidae ..... 55-60 

Astatoreochromis .... 165-1 77 

atlantica, Nidalia . . . . .109 

atlantica, Sagitta serradentata . . 234 



Cochlearius . 
contortus, Bulinus . 
cornigera, Aplysia . 
eronus, Haplochromis 
crunullae, Aplysia . 
cryptodon, Haplochromis 



55-62 

J 7 

302-304, 384 

I8O-1 82, 204 

304-3O6, 384 

97, 198-2oo, 204 



dactylomela, Aplysia 

278-284, 307-3IO, 382, 384-385 
dautzenbergi, Bulinus . . . .19 

decipiens, Sagitta . . . 239-240, 251 

delli, Aplysia nigra . 367, 370, 388 

denisoni, Aplysia . . . 310-3 12, 3&5 

depilans, Aplysia . 271, 274, 275, 357~36o, 385 
Dolabellinae . . . . . .269 

Dolabriferinae . . . . .269 

draco, Pterosagitta . . 246~7, 251 

dulitensis, Callosciurus notatus . -259 

dura, Aplysia . 36O-363, 385 



Balaeniceps . 

Balaenoptera 

Bathynella 

Bellonella 

bicolor, Macropleurodus 

Biomphalaria 

bipes, Aplysia Juliana 

bipunctata, Sagitta 

bocagei, Bellonella 

bocagei, Cereopsis . 

boissyi, Biomphalaria 

brasiliana, Aplysia 

brunnea, Aplysia nigra 

Bulinus 



Cactogorgia . 

calif ornica, Aplysia 271, 

Callosciurus . 

cedrosensis, Aplysia 

Cereopsis 

cervina, Aplysia 

chilotes, Haplochromis . 

chromogynos, Haplochromis 

Ciconiidae 

cinerea, Bellonella . 

Circoniiformes 

clavatum, Metalcyonium 



51-71 

125-161 

43, 44 

IOI-113-I2I 

76, 78, 80, 83, 96 
17-19 

364, 365, 368, 387 

228, 229, 231, 251 

107 

109, in 

17-19 

297-299, 382, 383 

. 370, 388 

. 3-28 



101, 105, 106, 114 
277, 294-297, 383 
258-265 
354-357, 374 
109 

299-302, 382, 383 
207-212, 217 
211, 212-214 

54-7o 
112 

53-70 

IIO, III 



elegans, Sagitta 
Eleutherobia 
enflata, Sagitta 
erythraeus, Callosciurus 
euchlora, Aplysia . 
Eukrohnia 
extraordinaria, Aplysia 

fasciata, Aplysia 

Fasculariidae 

ferox, Sagitta 

finlaysoni, Callosciurus 

flavimanus, Callosciurus 

foliacea, Nidalia 

forskali, Bulinus 

fowleri, Eukrohnia 

Fregata 

Fregatae 

friderici, Sagitta 

geographica, Syphonota 

Gersemia 

gestri, Haplochromis 

gigantea, Aplysia 

globosa, Physopsis 

globosus, Bulinus . 



• 233 

112 
240-24 1, 251 

260 
272, 374-377, 385 
221-225, 248-250 
275, 312-315, 385 

275, 315-3i8, 382, 386 
. 118 
232-233 
260 
260 
112 
19 
248, 249-251 
55-7° 
55-69 
228-229, 251 



• 374, 39i 

no, 118 

173, 182 

319-320, 386 

. 18, 19, 22 

19-25 
globosus jousseaumei, Bulinus (Physopsis) 25 



406 



globosus ugandae, Bulinus 
Glyphotes 
gracilis, Aplysia 
grandiflora, Thesea 
granti, Haplochromis 
granulata, Bellonella 
granulata, Nidalia 



haematobium, Schistosoma 
hamata, Eukrohnia 
Haplochromis 76-97, 165-166, 
Helisoma 

hexaptera, Sagitta 
Hoplotilapia retrodens 

Ibis 

inca, Aplysia 

inflatus, Bulinus 

Isidora 

Itephitrus 



Jabiru . 

japonica, Eleutherobia 
jousseaumei, Bulinus 
Juliana, Aplysia . 280 



keraudreni, Aplysia 
Kochella 

Koonungidae 
Krohnitta . 221-222 
kurodai, Aplysia 



labiatus, Lobochilotes 
labrosus, Melanochromis 
Laplysia 

lepidops, Spelaeogriphus 
Leptoptilos . 
Lepus . 
Lernaea 

leucomus, Callosciurus 
leucomus, Prosciurillus 
lividus, Haplochromis 
lobiancoi, Aplysia . 
Siphonata 
lobochilotes, Haplochromis 
Lymnaea 
lyra, Sagitta . 

macrocephala, Sagitta 
Macropleurodus 
macrospina, Bellonella 
maculata, Aplysia . 
maxillaris, Haplochromis 182, 
maxima, Sagitta . 235, 

Melanochromis 
melanopterus, Haplochromis 
melanotis, Nannosciurus . 
Menetes 



283 



225- 



22 

257-258-266 

320-321, 386 

102, 117 

86-90 

106-107, 113 

107 

. 3. 19 

248-249, 251 

172-173, 179-2i8 

10, 18 

238-239, 251 

80 

55, 58 
321-322 

12 



[09, 



7 
in 



• 58 
112 

. 3-28 
363-369, 386-387 

322-325, 387 
. Ill 

43 
226, 247, 249, 251 
325-326 

. 211 
211 

• 274, 275 
33-41 
58, 62 

• 274 

• 274 
262-264 
263-265 
205-207 

• 294 

• 274, 315 
211 

4 
235-237, 251 

245-246, 251 
76, 78, 80, 83 
. 117 
326-328, 387 
189-192, 204-205 
237-238, 245, 251 
211 
166, 192-194 

• 263 
, 260 



Metalcyonium 






IIO, III 


microdon, Haplochromis 




2OO-203, 204 


Microsciurus . 






263-265 


Microsciurillus 






263, 264 


minutus, Myosciurus 






. 263 


Monodella 






41,42 


morifera, Nidalia . 






. 112 


morio, Aplysia 


275 


328-3 


mouhoti, Aplysia . 






374. 391 


murinus, Prosciurillus 






262-265 


Myosciurus . 






261-265 


Nannosciurus 


260-262 


, 264-265 


Neaplysia . . 271 


273. 


294 


-297, 380 


neglecta, Sagitta 






245. 251 


Nephthyigorgia 






• 115 


Nidalia 




IOI 


-113-121 


nigra, Aplysia 


367, 369-370, 388 


nigrescens, Haplochromis 






194, 217 


nigrocincta, Aplysia parvula 






288, 291 


nigrovittatus, Callosciurus 






. 261 


nigrovittatus orestes, Callosciurus 




• 259 


Notarchinae . 






. 269 


notatus, Callosciurus 






258-264 



obesus, Haplochromis 

173, I82-183, 190-192, 204-205 
occidentalis, Astatoreochromis alluauda 174-177 
occidentalis, Nidalia 101, IO2-105, 113, 118 

oculifera, Aplysia . . 279, 332-334, 388 

owreae, Cercaria . . . . .239 



pacifica, Krohnitta 






• 247 


palustris, Lymnaea 




4 


Parabathynella 




• 43 


Paralemnalia 




. 118 


Paraxerus 




• 265 


parvidens, Haplochromis 






187 


, 194-198 


202-203, 204 


parvula, Aplysia 


271, 280 


287-291, 388 


paucidens, Haplochromis 




. 214 


Pelecani 




55-69 


Pelecaniformes 






53-7° 


Pelecanus 






55-7° 


peregra, Lymnaea 






4 


Pericarida 






44, 46 


Phaethon 






• 54 


Phalacrocorax 






55-70 


Phycophila . 




272, 273, 


374-377. 382 


physalus, Balaenoptera . 




125-i6i 


Physopsis 




3. 7-25 


plagiodon, Haplochromis 




205-2O7 


planctonis, Sagitta 


241, 


242-244, 251 


prevosti, Callosciurus 




. 260 


prodomus, Haplochromis 


79. 80, 


82-86, 88, 96 


Prosciurillus . 




261-265 


Pruvotaplysia . 271, 


273, 287- 


-294, 380, 381 


Pterosagitta . 




221-228, 


246-247, 251 









INDEX 






407 


pulchra, Sagitta 




• 245, 251 


Spelaeogriphacea . 




45. 46 


pulmonica, Aplysia 


335 


"337. 382, 389 


Spelaeogriphus 






31-46 


punctata, Aplysia 






spuria, Aplysia 






. 287 


271, 275, 279 


283, 284 


29I-294, 389 


studeri, Bellonella . 






112 


pusillus, Sciurillus . 




• 263 


studeri, Cereopsis . 
subtilis, Krohnitta 






in 
247, 251 


quoyana, Aplysia Juliana 




368, 387 


Sula 

sydneyensis, Aplysia 






55-7o 
348-350, 390 


rehderi, Aplysia 




337-340 


Syphonota . . 270, 


274. 


3i5, 381, 39i 


reticulata, Aplysia 






340-341, 389 








rex, Balaeniceps 






51 


Tamiscus 




• 265 


rigida, Bellonella 






112 


taurinus, Haplochromis . 




205 


rigida, Nidalia 






• 105 


tenuis, Sagitta 




229-231, 251 


robertsi, Aplysia 






342-343, 389 


Tethys .... 




• 274, 275 


robusta, Sagitta 






232-233, 251 


Thermosbaena 




42 


rubra, Bellonella 






112, 117 


Thermosbaenacea . 




41, 46 


Nidalia 






112 


Thesea .... 




102 








Tilapia 




187, 203 


sagamiana, Aplysia 




343-345, 382 


trivolvis, Helisoma 




10, 17 


Sagitta 


221 


-228-246, 251 


truncatus, Bulinus 




18 


sauvagei, Haplochromis 


76-8i, 85, 88, 211 


tryoniana, Aplysia pulmonica 




335. 389 


Schistosoma . 




• 3. 19 


Tullia .... 




271, 365 


Sciurillus 




261-264 








serratodentata, Sagitta 




234, 251 


vaccaria, Aplysia . 




371-374, 390 


sibogae, Aplysia Juliana 




368, 387 


vanderhorsti, Astatoreochromis 


166-167 


sibogae, Bellonella 




. 117 


variabilis, Bellonella 




112 


similis, Microsciurus 




. 263 


Rhodophytum 




. 112 


simpsoni, Cactogorgia 




. 106 


Varria . 271, 273, 


297- 


-354. 380, 381 


simus, Glyphotes . 




257-258-266 








Siphonogorgia 




115, 116, 117 


willcoxi, Aplysia 




350-352, 390 


Siphonota, see Syphonota 






winneba, Aplysia . 


352-354, 382, 390 


sorex, Aplysia 




• 363 








sowerbyi, Aplysia . 




345-347. 390 


xenognathus, Haplochromis 




. 90-95. 217 


speciosus, Itephitrus 




109, in 
















zetosios, Sagitta 




241 


242-245, 251 




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